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PROCEEDINGS OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


February 15, 1977 


Series 4, Volume 41, Number 1, 123 Pages, 45 figures 


THE OSTEOLOGY, CLASSIFICATION, AND RELATIONSHIPS 
OF THE EEL FAMILY OPHICHTHIDAE 


eo 
John E. McCosker 


Steinhart Aquarium, California Academy of Sciences 
San Francisco, California 94118 


CALIFORNIA ACADEMY OF SCIENCES/GOLDEN GATE PARK/SAN FRANCISCO 


February 15, 1977 


Series 4, Volume 41, Number 1, 123 Pages, 45 figures 


THE OSTEOLOGY, CLASSIFICATION, AND RELATIONSHIPS 
OF THE EEL FAMILY OPHICHTHIDAE 


By 
John E. McCosker 


Steinhart Aquarium, California Academy of Sciences 
San Francisco, California 94118 


ABSTRACT. A classification of the genera of the apodal family Ophichthidae is proposed on 
the basis of internal and external morphology, with particular emphasis on osteological 
characters, Specimens of 89 ophichthid species from 44 genera and comparative material 
from ten other apodal families were prepared for osteological examination, usually by a 
trypsin-based staining and clearing technique, and critically compared. Forty-nine ophichthid 
genera are recognized and are distributed among six tribes in two subfamilies. Diagnostic 
characters for the recognition of genera include the shape and condition of elements of the 
gill arch and hyoid arch, number and placement of branchiostegal rays along the hyoid, sus- 
pensorium elements, neurocrania, dentition, pectoral girdle elements, cephalic pore patterns, 
lateral line ossification, fin placement, and morphometric characters. The Ophichthidae are 
defined on the basis of their numerous overlapping branchiostegals, supraorbital canals 
united by a transverse commissure through the fused frontals, first and second epibranchial 
interconnections, absence of a palatine, and the separation of the pterygoid from the vomer. 
A monophyletic origin of the family from a congrid-like ancestor is proposed. An evolu- 
tionary history of the Ophichthidae is suggested, in which the subfamily Myrophinae has 
separated into two tribes and the subfamily Ophichthinae has radiated into four tribes. The 
validity of the family name Ophichthidae is discussed. The family names Ophisuridae, 
Myridae, Myrophidae, Muraenichthyidae, Echelidae, Neenchelidae, Aoteidae, Acanthen- 
chelyidae and Sphagebranchidae are synonyms of the name Ophichthidae. A comparison is 
made between an ophichthid classification based primarily on osteology and the previous 
classification, based primarily on external morphology. The results of two computer-pro- 
grammed classification schemes of species relationships within a single tribe are compared 
with a classification developed using traditional methodology. Alternate hypotheses are pro- 
posed to explain the log-normal inverse relationship between genera and the distribution of 
species among genera in the Ophichthidae. 


CALIFORNIA ACADEMY OF SCIENCES/GOLDEN GATE PARK/SAN FRANCISCO 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


TABLE OF CONTENTS Page 

LOST? OOTP [PUGS ae a a Ee oes ee 5 
LUST OE WBA Se an ee a ee ee Se ee eee ee 6 
PNGIIN ON IEEID GNA EINGTIS seat oe eee a ere ee ee A ees ee ee 7 
DONTUTRCC ET SATION i Sek ni on a Ro gn oe 9 
CGT rel etcetera re Ne ON RR a RPS 2 a ei mn eas Le et 9 
ISKOnVRO te © Piicehttni clan Glass iit Gell inp eeaea eee eEE 10 
Valli ditwaotastineratimilvadNalnn ees sons see ee ee Se Ses 10 
@steologicaleStuclesmotmthie © point Glivtl nical esses eee ee ee 11 
Fetal! SwAROMMNVinMNY Cn We (VOM AMM MCNS: csc ses pence aracnoeasaeiteleennassecnqeecereeenens 12 
SVMOPSISMm Olen © oli Cine en Glas SI fill Galt © meee eae 13 

VAT EAT at @) DB) S prereset eect ae eee cen kN dot AU cers. Seta we ee Lee 2 eens Neuere ae ee Ree 14 
AXON OMiLGe NIETO Seren 2: ae eee ee Seen ne te ee eee 14 
ING) ON RERITE NTI TVS mae se ee ip a eg al ak ee ie mere ae ree Ew Be Terre ee eee nr do 14 
INMare iia Summ sxcel I © Cl te sects cers re cor Sk nee nce Ran ae oS cee ne aE 115 
Statist calle NA Cth © CS pe ccseeretas ee. elec nee cece Se ee oe ee ence a eee 16 
OSTIEOLOGYAAN DOIN Gail @ INAS AINA GTI @ NAN eee ee ea 17 
INQUITROXCTITUI I eI Ns A ep ae Ae A es eR, SRT WR EB Oe tare SP ae etek ew OD 18 
SUSDEMSOnUin Mpa) Cala Stes eee eee ees oe eee ei ewe td eae ee tala eee ed eee 24 
(OVOVSTRCUNLETE SYSU TRS eee a Ea en SP RM oe CO ee te AY 26 
FLAVONIG!" AVC) OF IEE UI a a Ea REE ele Fa he ee a ne ee A Gee be ee ee ee 28 
GrTTBRPAN Gin Steet acta nee cere ne eke Ese ee SE ee we eet ne nd Pes ee SK NE 32 
Recto all Gite) Cie sere tek oom crete Bec Mee EN Ot ARG An ony ec ezcninrt se ceed ewes MAUR 33 

TRAN pall Same SV-S1CG tae see nce eaten Noe sec Se eee OLDEN Ee EP RCE NN ied SO SO ROE ee ER 36 
PSE) SIRS EXC CYS Bae ee tea a aS te Ee EN Re eh Ses eR erent 42 
(@alliclalleeS Kel ete ri ee ne ee cee ce eee ee eae ee eo 45 
NSCS TNT M CO NY ee a ee ee ee te eee 48 
TAUOINIOUNAN LBS ater AEE RE re ONE rns = ed AER Oe 48 
OsteolosicalsDetinitionmotmtiney. © fo lnt cht lnc ele een eee 49 
AnalviticalakeyatonthenGemetanot © pln htin cl ele eee seen eee en eens 50 

| EIUTOY Se GEA SH ST ae eS ee RES ONE en Oe eee Ne ee eer ne ee 56 
Subfamilial and Tribal Diagnoses and Generic Descriptions ........................---------------- 57 

S Ura fala Lame VAN @ PIMNIN a Cisse sete Ea cern cea eee cen ie aan See seed Sco Semen 57 
Tribe Benthenchelyini (Genera are listed alphabetically within each tribe) -....... 57, 

Tithe cae WANT POINT lies sees ee eee ie ead ee Re Se ect eine Ree Ve ee 57 
Sulfate @ pln clnthinaes<-cee.ccese ac cece ee oe cee en ers o ee Se cere ec ese ee 62 

al ftai to eee @all Hehe bn rps ae ee ee ee ee ee ee DE Eo 62 
inibemspiare bray Ghitiniee see ere te see tesa 2 ctor ee ie cael 2 eee 64 

Miriam aS Gel ml Glinitih i Id gaa ea ae ee eke Fe OF We cee ee Nt eee eee 70 

Wtloves CO fool pitied ni nin] Seen aes ene Bt ae eee eo LE ne ae ee Perea eee 73 
GompanisonmwithmeneviOUSn Glass Calti@ mSy esses eee eee eye 85 
ENWOILUNTIOUN (Ol TRIE COPRIICTRINRIIOYNE aoc ercnes ase ecneceenc ss Sesgee epee perpen Bop ee eS DO SEE EEE 85 
Relat Oms tio m CO mOthne ray A tne Ulli rrr Siemens eee eae era nnn 85 
EWOMUUNCO LH) AOMATUL MMM) tA COV TCT nyt CLS) ee 86 
MMVIROFOLMIID ine! TEMG MEMEIMITDN — co sececccenne conecee eee seece ener ecm eee Rs aeneaerecerer pRoDAD HOSS EEEOSECEEEBEES 88 
COWS RTSI LU MMUMLAN = ack eas es eer eee NO oe cer yt UC PCR er eee eee eee ete sks 89 
Sita ge bral clini iif eaten ee eee ene ete noo ee eee ee tee ER ree 91 

Beals earl GIG ip meee erecta meme nc tines ete eee Pi hy Chel Cle a cee eat Wee A he ad 94 
Gallechyel\itriiiameteen erate eee hes Nae Deer: oe a Td oo ea ee Be ete 94 
ZOOGEOGRAPHY AND COMMENTS ON OPHICHTHID SPECIATION ..............-.--...--------- 96 
[LUPTTELRSVN TIRE (CTE Yo rey a AS ace aE ae Sa sad cn ER SR Ie a, ue eee oe, 101 
AB KES eee ete ir, ener A RG Ser ee oe Ao mW ee a eee Har ek ee Sea 108 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Figure 


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HRHPA HR PWWWW WWW WwW Ww 
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LIST OF FIGURES 


An Early Illustration of an Ophichthid, presumably Ophisurus serpens .................- 
NedrocraniUimmotm © polaiclatlnusweZ Ojo no Gliliga seen eee en 
NetnocraniUnn nots © a hniGiathtiSiez Op lO Ginliigesesae senses tenner 
Neuro Granite Bem thiernGin@ isin Gel itil irene eae eee ne en cE ES 
NG unOGraniUinmn iO fil VV{i@ Toil Sav cli ames ees En oo 
Neuro cranium One Meira eniliGithnySirGinilenmS 1S. esssssee teen ence en ee ee 
NeurocranUmmotm @alll@Gineliysiiimalrinp © nali Us apenas eee ane nee ee ee eee 
Neuro craniUinmoteaGiitinyaoUrsies eC] a\Gii@ [a Simsesseeere eee arenes eee 
NeURocAniuinn @ir SWiGiodminWS [OOMMIUS ccecee eer cece eee eee rsa eeeeeSeeeeee 
INEUKOcraniUMmOneBascaniGhth/Sm palmar C SiS esse tes tes scee eee eames ae 
NGUROGMIUIIN Cir IMAM AIS: SSVSTRUITWIS xcesseeccese cree ese c cence eeee coe cer eno a cea eeE Ree Re eem = HES aon nEnee 
@xoliths of Several’ @phichtinid Species EA ae Boe rin Se See 
Fleadmskeletommons © piGhitinUsiez OO © Cilia eeeeeea eee ee neon E 
Suspensonum) and jawsnot Ophichthus Zophochiin sss 
RightsPostonbital/ Series) of BrachySOmoOpinis) SAUMOPSIS, cess eee ee een 
Maxilla-VomensApposition: of Severall @yolniehnthyics wes cee eee 
Hyoid Arch and Branchiostegals of Ophichthus zophochir, an Ophichthine, 

ANG eel ea Gitay Sin Gln C1aSTSPaclan NAW/© [Ohi Cleese eee ec 
GillArcheskeletonmotm© piiiGitinustez.o in @ Cin aes ene ee 
Pectoral Girdle of Various Representative Ophichthines .................--------------0---00--- 
Pectoral Girdle of Various Representative Myrophines .............-..------:---------0000100+++= 
GephaliesRateralissSystembpanGeASSOGlatec | BOnes messes eee 
Lateral Line Ossicles of Representative Ophichthines .......................2.-.0--00ecceeeeeeeeeeeee 
Lateral Line Ossicles of Representative Myrophines .................-.-.--------------------------+ 


Cephalic Pore and Surface Sensory Papillae Development in Two Ophichthids .... 
Anteriormost Five Vertebrae of the Type Genera of the Tribes of Ophichthids .... 
Trunk and Caudal Vertebrae of Ophichhtus zophochir ...................-.--.-..2200--+-------+- 
Caudal Skeleton of Ophichthus zophochir ................-.......------- sate 5, SONIA ei et Oe 
GaicaleSkeletonmo te Vino phils aval Cymer 
Comparative Anatomy of Congrid and Ophichthid Digestive Tract and 

(GES. “BB V0 Kol eir Re ee ok A a, a as ae emetic a ek dill pe gee 41 hw Wn BO re aa 


Vomer, Maxillae, and Pterygoid of Ahlia egmontis and Myrophis vafer ................ 
Diagrammatic Representation of a Species with Well Developed Head Pores ...... 
Representation of Underside of Callechelyin Smouts .................22.--200000-222222000e--eee-e= 
Diagrammatic Representation of Posterior Trunk Vertebrae ................220.-00.---------- 
Diagrammatic Representation of Head and Pectoral Fins of Two Ophichthins ...... 
Diagrammatic Representation of an Ophichthin with a Fringed Upper Lip —.......... 
@©penculaneSenlesoteMiyropmismVvaret ss. = eee ee en Pe ee 
Proposed Evolutionary Relationships of Ophichthid Tribes —.................0222000.-...---- 
PROPOSE MEV. UETO MO itiatin Cal VAN TO [olin ele eee eee Nn 
RrOPOSEEEVO]UTO MMO Tate © knit Ghat nna eee ee ae ee sue en pS 
BLODOSEASEVOUTLONMOfatheMs Pliage ali Glniliiimeesemees seers eee 
BrODOSCGMEVOUTONMO fe tiveN Bas Gali Glitnir eres eee 
PROPOSCU MEV OUT ONMmOl mtn em alll Ec lne lly, iinet eee 
Phenogram of the Relationships of the Callechelyini, Using Program WVGM ...... 


Interrelationships of Species Groups of the Tribe Callechelyini, as Defined by 
PORN LEO © OI ES cee ne Sen salar Sean re Oe MN MeN RINE NTS ay eee eeaito 
Distribution of Species Among the Genera of the Ophichthidae, 

Gobiesocidae and _ Salariini 


6 


SERIES 4, V.41, #1 McCOSKER — EELS 


LIST OF TABLES 


Table Page 
PD eMtitOnmOimthem Gem enaeotin© plat cliche epseceee ee ease eee tenes eee snes 108 
2 Number and Location of Branchiostegal Rays of the Species of the Ophichthidae .... 109 
See GilleArcheGonelitio nny th yey © po ai ltl rea Crees eee etme ns enn seen 110 
aL Gill “Aely Geommvaliittiormy flim: me IMAGO Son cere eee ee CeCe 111 
5 Lateral Line and Cephalic Pore Conditions in Ophichthine Genera and Subgenera .. 112 
6 Wereorll Coume Or Waltouws Wyolnichntimicl SlxXheS .sceesecs access cage ceccenennonccosenseseeseons Ms 
7. ‘Gharacteristics of the Ophichthidae and Related Eel Families... 116 
8 Morphological and Meristic Characters of the Species of the Callechelyini —........... 7, 
9 Characteristics of the Species of Callechelyini Used in Programs REGROUP 

PATI) GR VIN Gt ANS ose ta steel NC 2 Sr ec SA Sao J we nyu ee Se, Seen ne 118 

10. Wistribution“of Certain Ophichthid) Gemena: .....22. 2 cen2-05s2-cece ace oe cece 119 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


ACKNOWLEDGMENTS 


The majority of this work is from my doctoral 
dissertation done at the Scripps Institution of 
Oceanography, University of California, San 
Diego, under the direction of Richard H. Rosen- 
blatt and Carl L. Hubbs. | sincerely thank Profes- 
sor Hubbs for his advice and guidance through 
the capricious nuances of zoological nomencla- 
ture, and Professor Rosenblatt for his invaluable 
advice, encouragement, and patience throughout 
the duration of my graduate studies. 

| wish to express my gratitude to the following 
individuals who have made various specimens 
available: Marie-Louise Bauchot, Paris Museum; 
Jacques Blache, Centre ORSTOM,; James E. Bohlke, 
Academy of Natural Sciences of Philadelphia; 
Peter H. J. Castle, Victoria University of Welling- 
ton, New Zealand; Lev Fishelson, Hebrew Uni- 
versity; John E. Fitch, California Department of 
Fish and Game; Warren C. Freihofer, then of 
Stanford University; Robert H. Gibbs, Jr. and 
Robert H. Kanazawa, National Museum of Natural 
History; William A. Gosline, then of University of 
Hawaii; Naercio A. Menezes, Universidade de 
Sao Paulo; Hans Nijssen, Zodlogisch Museum 
Amsterdam; John R. Paxton and Douglass F. 
Hoese, Australian Museum; John E. Randall, 


Bernice P. Bishop Museum; Tyson R. Roberts, 
Museum of Comparative Zoology, Harvard Uni- 
versity; C. Richard Robins, University of Miami 
Marine Laboratory; Margaret M. Smith, Rhodes 
University; Enrico Tortonese, Museo Civico di 
Storia Naturale, Genova; Boyd W. Walker and 
John Bleck, University of California, Los Angeles. 
| am particularly grateful to William N. Esch- 
meyer and the staff of the California Academy 
of Sciences for making the extensive and critical 
material from the George Vanderbilt Collections 
available to me. 

Thanks are also due to the following individ- 
uals: Ira Rubinoff and the staff of the Smithsonian 
Tropical Research Institute for assistance during 
my tenure as a Smithsonian pre-doctoral research 
fellow; Edward W. Fager and John H. Wormuth 
for assistance with computer programs; students 
and colleagues at Scripps Institution, and in par- 
ticular Joseph F. Copp and Donald M. Dockins, 
for aiding in numerous ways; Richard H. Rosen- 
blatt, Carl L. Hubbs, Robert R. Hessler, and Peter 
Paul Vaughn for their critical reading of my dis- 
sertation; and my wife, Sandra, for her help and 
encouragement. 


SERIES 4, V.41, #1 McCOSKER — EELS 


SERPENT MARIN. 


UMMA IFEOES 7570, i 
4 VILL —oe 
wo MCU TET poe 
Ui eS: 
/ ye tH LCC Gay, Sk < 
RAY 1 "ag, —\ IE eS DL 
uyetae 4 = gat nay 
sete Se SR 
Lhe. cAS-. ‘J S % 
” Lo AS 5 2 
% WR J, = 
P ; 4 SR 2 3 
W pp ee , wee Zz E 
Wir rss ‘WS if? E 
oe, A > ince s os 
VAM \S= OB Ps 
1 Fy Wi If gE 5 =.= Zé ORT) 
aN IX: = Aa 
Bs : 5 A 
z N ~ 2 
° as 7 Se ; 
\ ye £[F 
aN LE 
AG * Za w= BS 
Af, e Ffg= 
\\ = 
FX 
SS 
— 
' {! 1, 4, 
serpens, 


An early illustration of an ophichthid eel, presumably Ophisurus 


Figure 1. 
from Mattioli’s Commentaires (1568). 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


THE OSTEOLOGY, CLASSIFICATION, AND RELATIONSHIPS 
OF THE EEL FAMILY OPHICHTHIDAE 


John E. McCosker 


INTRODUCTION 


General 


The Ophichthidae comprise a large family of 
mostly fossorial eels limited to continental shelf 
depths of all tropical and subtropical oceans. The 
Ophichthidae contains more than 200 species 
here distributed among 49 genera, representing 
perhaps the greatest diversity of anatomical spe- 
cializations within a single apodal family. The 
resemblance of many ophichthids to snakes has 
earned them the common name of ‘‘snake-eels,’” 
and along with the morays, have evoked many 
sea-serpent legends among tropical coastal peo- 
ples. Perhaps the earliest illustration identifiable 
as an ophichthid (fig. 1) was that of Mattioli 
(1568) p. 388, probably based on an adult Ophi- 
surus serpens. The first described ophichthid 
species, Muraena ophis, was the second apodal 
fish in Linnaeus’ Systema Naturae (1758). 


The taxonomic treatment of the Ophichthidae, 
and of apodal fishes in general, has been in con- 
stant flux since the eighteenth century, and only 
within the past two decades has a coherent con- 
cept of the Ophichthidae become realized. Pre- 
vious theories of the interrelationships of ophich- 
thid genera have been based on such trivial char- 
acters as fin position, dentition types and colora- 
tion. It is now generally accepted that the osteol- 
ogy more conservatively reflects the phylogeny 
of a group and is less affected than external 
morphology by minor evolutionary adaptations. 
The unsatisfactory state of the classification of 
the ophichthids has been recognized by numer- 
ous authors (Myers and Storey, 1939; Myers and 
Wade, 1941; Gosline, 1951a; Smith, 1964; Rosen- 
blatt and McCosker, 1970; Robins and Robins, 
1971; Castle, 1972) most of whom have con- 
cluded that an osteological revision of all in- 
cluded genera was necessary for a proper under- 
standing of the family. The aim and scope of this 
study have been directed to that objective. 


10 


History of Ophichthid Classification 


The 218 years since Linnaeus’ (1758) descrip- 
tion of the first ophichthid species have witnessed 
a plethora of generic and specific names applied 
to the Ophichthidae. The first generic name 
properly applied to an ophichthid was Ophich- 
thus (Ahl, 1789), which should more properly 
have been written “Ophichthys’”. The emenda- 
tions of Ahl’s generic spelling by subsequent 
authors have resulted in confusion over the gen- 
eric and familial names. This problem was briefly 
treated by Gosline (1951a: p. 298) and is further 
illuminated in the following discussion. 


The treatment of apodal taxonomy was in con- 
stant flux during the eighteenth and nineteenth 
centuries, which accounts for much of the lack of 
uniformity in nomenclature. Swainson’s (1838) 
treatment of the eel-like fishes was based largely 
on Cuvier’s work (1817), and was the first at- 
tempt at an arrangement of the eels into family 
groups. Swainson (p. 215) designated the ‘“Mur- 
aenidae (as) having two branchial spiracles in 
their ordinary position, and the Sphagebranch- 
idae, or sea eels, where the branchial spiracles 
are either close together or united into one.” 
Confusion ensues on the following page where 
the family name Gymnarchidae is apparently con- 
sidered synonymous with the Symbranchidae, 
and further evidenced in his discussion (p. 218) of 
the gill openings, by his statement that ‘‘among 
the Gymnarchidae, or sea eels, for instance, they 
are close together and united under the throat as 
in Sphagebranchus.”” Swainson divided the Mur- 
aenidae into two subfamilies, the Anguillinae and 
the Muraeninae, both of which contained species 
now known to be ophichthids. McClelland (1844) 
realigned the apodal classifications of Swainson 
and Cuvier and created the family Ophisuridae 
to include eels with a rayless caudal contain- 
ing the genera Leptognathus (=Ophisurus), Ophi- 
surus, and Ophithorax (=Ophichthus). Kaup 
(1856a,b) disregarded most of McClelland’s 
classification without comment, but did retain 
the name Ophisuridae. Kaup divided the apodal 
fishes into two ‘sections’. These were the Cryp- 
tomycteres (containing only the Ophisuridae) 
which included those eels with labial nostrils, and 
the Phaneromycteres which contained all other 
apodal families. The ophisurids were divided into 
three subfamilies, the Ophisurinae, the Sphage- 
branchinae, and the Myrophinae (containing 
Myrus, Myrophis, and Muraenichthys). Bleeker, 
in his Systema Muraenorum Revisum (1865), rec- 
ognized the family Ophisuroidei and considered 


SERIES 4, V. 41, #1 McCOSKER — EELS 


the Myrophinae (as Myriformes) to be a subfamily 
of the family Congroidei. Ginther (1870) con- 
siderably revised previous classification by plac- 
ing the majority of the known eels into a single 
family, the Muraenidae, which he divided into 
ten ‘‘Groups’’. The Ophisuridae of earlier authors 
was divided into two groups, the Ophichthyina 
containing those species with a rayless caudal 
[comprising the genera Liuranus (sic) with a 
single species and Ophichthys with at least 78 
species], and the Myrina containing those with a 
rayed caudal (comprising Myrus, Myrophis, Para- 
myrus, Chilorhinus, and Muraenichthys). Gwtn- 
ther’s groups were elevated to family rank by 
Jordan and Davis (1891). The Myrinae became 
the family Echelidae (Jordan and Davis considered 
Myrus a synonym of Echelus) and the Ophich- 
thyina of Giinther (actually, the Ophisuroidei of 
Bleeker) became the Ophisuridae. Uncertainty 
concerning the synonymy of Myrus and Echelus 
resulted in the changing of the name Echelidae to 
Myridae by Jordan and Evermann (1896) and by 
Jordan and Snyder (1901). The family name 
Ophichthyidae, derived from Gunther's Ophich- 
thyina, first appeared in Jordan and Evermann 
(1896). The authors rejected the name Ophisuri- 
dae and considered Ophisurus a synonym of 
Ophichthus. 


Validity of the Family Name 


Confusion relating to the spelling of the 
Ophichthyidae with a “y’’ relates to the correc- 
tion by earlier authors of Ahl’s (1789) spelling of 
Ophichthus. The generic name is from the Greek 

re PF , meaning snake, and ixvovys, mean- 
ing fish, and would more correctly have been 
written Ophichthys. Bleeker, Gunther, and other 
classicists emended Ahl’s generic spelling (see 
Jordan and Gilbert, 1882), but Jordan and his 
later co-authors returned to Ahl’s original spell- 
ing. The retention of the original spelling of the 
generic name but the usage of the emended fam- 
ily name is illogical, and according to the Inter- 
national Code of Zoological Nomenclature, in- 
correct, as Gosline (1951a) has pointed out. Most 
recent authors, with few exceptions, have used 
“Ophichthidae”’ and ‘“Ophichthus”. 

A serious difficulty however exists, concerning 
the earlier family names proposed by Swainson 
and by McClelland. The inconsistencies in Swain- 
son’s usage of Sphagebranchidae would invalidate 
it as a family name. Although Sphagebranchus 
Bloch (1795) was then a valid genus and properly 
an ophichthid, Gymnarchus, a gymnarchid, was 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


later shown not to be an apodal fish. The prob- 
lem of recognizing McClelland’s usage of Ophi- 
suridae was avoided by Gosline who stated that 
“whether or not Ophisurus is a valid genus is a 
moot nomenclatorial question. Consequently | 
prefer not to use for this family, at the present 
time, the little-known and possibly invalid name 
Ophisuridae.”’ The logical solution to this nomen- 
clatural dilemma seems to be the invoking of the 
plenary powers of the International Commission 
on Zoological Nomenclature. The suppression of 
the rarely used name Ophisuridae in favor of the 
universally recognized name Ophichthidae would 
clearly be in the interests of stability. 


Osteological Studies of the Ophichthidae 


Regan (1912) was the first to attempt an osteo- 
logical definition of the Ophichthidae. Most im- 
portantly, his emphasis on the fused frontals of 
the ophichthids, congrids, and relatives has re- 
mained as a fundamental character in our con- 
cept of eel evolution. The other osteologicai 
characters identified by Regan (‘‘caudal vertebrae 
with transverse processes” and ‘‘maxillaries articu- 
lating with ethmoid near the end of the snout’) 
only described certain members of the family. His 
separation of the congrids from the ophichthids, 
on the basis of their long and slender rather than 
vestigial neural spines, was also an important ob- 
servation. Trewavas’ (1932) apodal classification 
scheme followed Regan’s characterization of the 
ophichthids. For nearly two decades, subsequent 
ophichthid studies dealt only with superficial 
characters. An exception was that of Myers and 
Storey (1939) who noticed the overlapping of the 
branchiostegal rays in ophichthid species. They 
pointed out that these rays are similar to the 
“jugostegalia’’ described by Parr (1930) in echelid 
eels. On that basis, and other external morpho- 
logical similarities, Myers and Storey suggested 
that the Echelidae might be merged with the 
Ophichthidae. 

Gosline (1950, 1951a, 1951b, 1952), in a series 
of papers, analyzed the species referred to the 
Echelidae and the Ophichthidae. His osteological 
study of Kaupichthys diodontus (1950) demon- 
strated that its osteology precluded its inclusion 
in the same family with Muraenichthys cookei. 
He found (p. 312-314) that K. diodontus differs 
in having sutured frontals and non-overlapping 
branchiostegal rays, as well as several other char- 
acters which later proved to be non-definitive. 
Lacking a specimen of Echelus myrus, Gosline 
was unable to further define the Echelidae, but 
suggested that species of Myrophis and Muraen- 


11 


ichthys might be referred to the Ophichthidae, to 
comprise the subfamily Myrophinae, on the basis 
of their “basket-like arrangement of the numer- 
ous, long branchiostegal rays.” In a following 
paper, Gosline (1951b) described the osteology 
of Chilorhinus brocki (=C. platyrhynchus) and 
related it to Kaupichthys diodontus. He suggested 
that the external similarities of species of Kaup- 
ichthys, Chilorhinus and ophichthids are ‘‘the re- 
sult of parallel evolution and not of close genetic 
relationship.” His prediction that Echelus, once 
examined on an osteological basis, would prove 
to be confamilial with Kaupichthys was incorrect 
(Gosline, 1952; Bohl ke, 1956a). 

Gosline (1951a), in a more comprehensive 
paper, prepared the first diagnostic treatment of 
the Ophichthidae. His study, however, was limi- 
ted to those species occurring in the Hawaiian 
area and thus did not include several critical 
genera. He compared the Ophichthidae with the 
Congridae (primarily Conger), and concluded that 
the ophichthid conditions are derived from, and 
more advanced than, those of their more primi- 
tive congrid ancestors. Two subfamilies within 
the Ophichthidae were recognized (the Myro- 
phinae and the Ophichthinae) although osteo- 
logical differences other than the caudal skeleton 
were not defined. In a subsequent paper, Gosline 
(1952) described the morphology of Echelus 
myrus in detail and concluded that it was refer- 
able to the ophichthid subfamily which contained 
Myrophis and Muraenichthys. On that basis he 
changed the subfamilial name of the Myrophinae 
to Echelinae. The results of that study did not 
alter his earlier (1951a) diagnosis of the family. 

Subsequent studies dealing with ophichthid 
osteology followed Gosline’s (1951a) general defi- 
nition of the family, but amended his diagnosis 
to include genera that he had not examined. 
Bohlke (1960) added Pseudomyrophis, and provi- 
sionally Neenchelys, to the Ophichthidae.In do- 
ing so, he expanded the familial diagnosis to 
allow the following: posterior nostrils either lat- 
eral or labial; maxillary articulation variable in 
position along the ethmoid; pharyngeal openings 
of the branchial clefts may be reduced; trans- 
verse processes of the caudal vertebrae either 
present or absent. 


Nelson’s (1966a) analysis of apodal gill arch 
conditions found most ophichthids to be ‘‘dis- 
tinguished in having the proximal ends of the 
dorsal parts of the first and second arches con- 
nected through a continuous cartilage, a peculiar- 
ity not present in any other of the eel families 
studied.’’ Certain generic lineages were identified 


12 


on the basis of their gill arch configurations. Nel- 
son's (1966b) study of the osteology of Neen- 
chelys buitendijki confirmed Bohlke’s earlier sup- 
position of its placement within the Ophichthidae. 
Nelson separated the ophichthids from the con- 
grids in the following manner: posterior nostril 
usually opening on the ventral surface of the 
upper lip; tongue adnate; branchiostegal rays 
overlapping along the midventral line; supraorbi- 
tal canals united by the transverse frontal com- 
missure; neural spines absent. Castle’s (1972) 
osteological study of Benthenchelys cartieri sum- 
marized the diagnoses from Gosline’s, BOhlke’s, 
and Nelson’s earlier works, but did not contribute 
to or amend their diagnoses. 


Familial Synonymy of the Ophichthidae 


The Ophichthidae, as currently recognized, in- 
cludes several families which were until recently 
considered distinct. The basis and validity of 
studies resulting in these actions are discussed 
below. 

The family Neenchelidae was erected by 
Bamber (1915) to contain Neenchelys microtretus, 
a new genus and species from the Red Sea. It 
was considered to be closely related to the Mur- 
aenesocidae as defined by Regan (1912). A sec- 
ond neenchelid, N. buitendijki, was described by 
Weber and de Beaufort (1916) from the Indo- 
Australian archipelago. The family received no 
further definitive treatment until Bohlke (1960), 
on the basis of Bamber’s description, suggested 
that Neenchelys may be related to species of 
Pseudomyrophis and thus properly be considered 
an ophichthid of the subfamily Echelinae (sensu 
Gosline, 1951a, 1952). The collection of adequate 
material of N. buitendijki and its osteological ex- 
amination by Nelson (1966b) supported Bohlke’s 
prediction. Nelson’s (1967) examination of the 
holotype of N. microtretus confirmed the recog- 
nition of the Neenchelidae as ophichthids in the 
subfamily Echelinae (herein considered as Myro- 
phinae). 

The Acanthenchelyidae also belongs in the 
Ophichthidae. Family recognition was short-lived, 
consisting of Jordan, Evermann, and Clark’s (1930) 
elevation of Acanthenchelys Norman. (errone- 
ously attributed to Regan) to family status, but 
this was largely ignored by later authors. Randall 
and Robins (1966) relegated Acanthenchelys to 
the synonymy of Ophichthus, an action which is 
followed here. 

The Aoteidae are provisionally included in the 
Ophichthidae. The Aoteidae were first recognized 


SERIES 4, V.41, #1 McCOSKER — EELS 


as ophichthids by Castle (1967), who referred the 
single aoteid species to the genus Muraenichthys. 

The family Echelidae (=Myridae, Myrophinae, 
Myrophidae, and Muraenichthyidae), has been a 
catch-all group with a checkered history. The 
echelids were considered congrid or muraene- 
socid relatives by most nineteenth century au- 
thors. Bleeker (1865), for example, considered 
the Myriformes (containing Myrophis, Echelys, 
and Muraenichthys) to be a subfamily of the 
Congroidei. Kaup (1856a,b) was exceptional in 
placing considerable importance on the labial 
nostril condition, and in allying the Myrophinae 
with the Ophisurinae as a single unit which ex- 
cluded the congroids and relatives. Twentieth 
century authors considerably expanded the Myro- 
phidae (which was to become known as the 
Echelidae, fide Jordan and Evermann, 1896) to 
include as many as 22 genera at various times 
(Schultz and Woods, 1949). The dissection of this 
large and cumbersome family was initiated by 
Myers and Storey (1939), and was followed by 
Schultz and Woods (1949) and by Gosline (1950, 
1951a, b, 1952). Myers and Storey noted the pres- 
ence of accessory branchiostegal rays (the “‘jugo- 
stegalia’’ of Parr, 1930) in both ophichthids and 
echelids, but were hesitant to merge the families 
without an extensive anatomical examination. 
Gosline (1951a) established the similarities of the 
two families on an osteological basis and included 
the genera Myrophis and Muraenichthys in the 
Ophichthidae to comprise the subfamily Myro- 
phinae. His subsequent (1952) osteological ex- 
amination of Echelus myrus resulted in its inclu- 
sion into the family, and the replacement of the 
name Myrophinae with Echelinae. The present 
study has demonstrated that Gosline was correct 
in considering the Myrophines to be ophichthids, 
but erred in including Echelus with the Myro- 
phinae. Although certain authors have continued 
to recognize a family Echelidae for the forms 
with caudal rays (e.g., J. L. B. Smith, 1962; Blache, 
1968), no convincing arguments have been pro- 
posed which would merit familial separation. 
Blache’s (1968: 1501) continued usage of the 
name Echelidae, with the justification that 
“.,.nous sommes également, tout a fait, partisan 
de cette position et nous ne conservons ici, la 
famille des Echelidae, que pour des raisons arti- 
ficielles de commodité taxonomique,” is both 
illogical and incorrect. 

Further substantiation of the inclusion of the 
Echelidae with the Ophichthidae is evidenced in 
the leptocephalus larval stage. (The leptoce- 
phalus of Neenchelys has not been identified.) 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Eel leptocephali display evolutionarily conserva- 
tive features that could prove useful in phylogen- 
etic investigations (Castle, 1965, 1967), yet the 
problem of generic and specific identification 
still remains and has precluded their usage in 
this study. It is important to note, however, that 
the morphology of ophichthid leptocephali re- 
flects the family grouping fairly well. Castle (1965) 
and D. Smith (unpub. MS) have diagnosed the 
ophichthid leptocephalus as moderately elongate 
when full grown, characteristically possessing gut 
thickenings or loops which usually accompany 
swellings of the pronephric ducts, conspicuously 
pigmented with patches of chromatophores which 
occasionally occur on the head, at various points 
along the gut, and often on the myosepta, lateral 
caudal midline, and dorsal and anal bases, and 
possessing a moderate to blunt tail. Castle (1965: 
98) has stated that ‘‘the pectoral remains obvious 
throughout larval life, probably even in those 
ophichthids which show loss of the pectoral in 
the juvenile and adult.’’ Phylogenetic implications 
at the generic level would therefore be further 
evidenced in this conservative larval condition if 
Castle’s suggestion proves correct. For example, 
his tentative identification of Leptocephalus Mur- 
aenichthys sp. is based on a leptocephalus pos- 
sessing a short, rounded pectoral fin (Castle, 
1965: figs. 2F, G), not unlike that of Myrophis 
(Eldred, 1966; Castle, 1965: figs. 3e-f). 

In contrast to the above mentioned families, 
the Macrocephenchelyidae was incorrectly synony- 
mized with the Ophichthidae. This family, known 
only from the holotype and damaged paratype 
of Macrocephenchelys brachialis Fowler, was syn- 
onymized without comment with the Ophichthi- 
dae by McAllister (1968: 85). Robins and Robins 
(1971) have re-erected the family on the basis of 
a thorough osteological examination of the para- 
type. They have shown its affinities to be with 
the Congridae and referred it to the superfamily 
Congroidea. Macrocephenchelys displays several 
characters quite divergent from the Ophichthidae, 
including the extensive ossification of the bran- 
chial apparatus (yet there is no lower pharyngeal 
tooth plate), the absence of the transverse frontal 
commissure of the cephalic lateralis system, and 
the presence of eight stout branchiostegal rays 
and a complete palatopterygoid arch. 


Synopsis of Ophichthid Classification 


The listing of nominal taxa below summarizes 
the taxonomic conclusions of this study. Full de- 
scriptions of new taxa and complete generic syn- 
Onymies are presented later in this paper. 


13 


Subfamily Myrophinae 
Tribe Benthenchelyini 
Benthenchelys Fowler 1934 
Tribe Myrophini 
Ahlia Jordan and Davis 1891 
Muraenichthys Bleeker 1853 
Subgenus Muraenichthys Bleeker 1853 
Subgenus Scolecenchelys Ogilby 1897 
Myrophis Lutken 1851 
Neenchelys Bamber 1915 
Pseudomyrophis Wade 1946 
Schismorhynchus McCosker 1970 
Schultzidia Gosline 1951 


Subfamily Ophichthinae 


Tribe Callechelyini 
Aprognathodon Bohlke 1966 
Callechelys Kaup 1856 
Letharchus Goode and Bean 1882 
Leuropharus Rosenblatt and McCosker 1970 
Paraletharchus McCosker 1974 


Tribe Sphagebranchini 

Achirophichthys Bleeker 1865 

Apterichtus Duméril 1806 

Caecula Vahl 1794 

Cirricaecula Schultz 1953 

Hemerorhinus Weber and de Beaufort 
1916, incertae sedis 

Ichthyapus de Barneville 1847 

Lamnostoma Kaup 1856 

Stictorhinus Bohlke and McCosker 1975 

Yirrkala Whitley 1940 


Tribe Bascanichthyini 
Allips McCosker 1972 
Bascanichthys Jordan and Davis 1891 
Caralophia Bohlke 1955 
Dalophis Rafinesque 1810 
Ethadophis Rosenblatt and McCosker 1970 
Gordiichthys Jordan and Davis 1891 
Leptenchelys Myers and Wade 1941 
Phaenomonas Myers and Wade 1941 


Tribe Ophichthini 

Aplatophis Bohlke 1956 

Brachysomophis Kaup 1856 

Cirrhimuraena Kaup 1856 
Subgenus Cirrhimuraena Kaup 1856 
Subgenus Jenkinsiella Jordan and 

Evermann 1905 

Echelus Rafinesque 1810 

Echiophis Kaup 1856 

Elapsopis Kaup 1856 

Evips McCosker 1972 

Leiuranus Bleeker 1853 


14 


Malvoliophis Whitley 1934 
Myrichthys Girard 1859 
Mystriophis Kaup 1856 
Ophichthus Ahl 1789 
Subgenus Ophichthus Ahl 1789 
Subgenus Microdonophis Kaup 1856 
Subgenus Centrurophis Kaup 1856 
Subgenus Coecilophis Kaup 1856 
Ophisurus Lacépéde 1800 
Phyllophichthus Gosline 1951 
Pisodonophis Kaup 1856 
Pogonophis Myers and Wade 1941 
Quassiremus Jordan and Davis 1891 
Scytalichthys Jordan and Davis 1891 
Xyrias Jordan and Snyder 1901 


METHODS 
Taxonomic Methods 


Osteological examinations, whenever possible, 
were based on entire stained and cleared speci- 
mens. Rare specimens and holotypes were studied 
by gill arch removal and radiographic examina- 
tion. Radiographs were prepared using a General 
Electric 40 KV x-ray unit and Kodak Industrial 
Type M film. Radiographs were either examined 
under a dissecting microscope or from photo- 
graphic enlargements. Stained and cleared gill 
arches or entire specimens were prepared using 
the trypsin-preparation method of Taylor (1967) 
and, in certain instances, the modifications of 
Miller and Landingham (1969). Neurocrania were 
prepared by dissection, soaking in a 5-7 percent 
potassium hydroxide (KOH) solution to remove 
the flesh, and staining in an alizirin bath. Sutures 
along the dry skulls became more apparent dur- 
ing examination when painted with pure glycerin 
using a fine camel’s hair paint brush. Certain 
skulls were disarticulated in a 7-10 percent KOH 
solution to better identify certain sutures. Draw- 
ings were made using a camera lucida attach- 
ment on a Wild dissecting microscope. 


Gill arch terminology is that of Nelson (1969). 
Bone terminology follows that of Asano (1962) 
with certain modifications that are identified in 
the section dealing with bone complexes. 

The following measurements, used in the gen- 
eric key and descriptions, are defined as follows: 

Head length. Measured from the snout tip to 
the posterodorsal point of the gill opening. 

Trunk length. Measured from the posterodorsal 
point of the gill opening to mid-anus. 

Tail length. Measured from mid-anus to the tail 
tip. 


SERIES 4, V.41,#1 McCOSKER — EELS 


Inclination of the suspensorium. The suspen- 
sorium is considered to be ‘anteriorly inclined’’ 
if the angle formed by the midlines of the hyo- 
mandibular and the mandible (when the mouth 
is closed) is greater than 90°. If the angle is less 
than 90° the suspensorium is considered to be 
“nosteriorly inclined’. This measurement is some- 
what subjective, and made either from radio- 
graphs or observations of stained and cleared 
specimens under the dissecting microscope. 

All fish lengths are listed as total lengths. Gen- 
eric descriptions and diagnoses were based on 
adults unless otherwise stated. 


ABBREVIATIONS 
Anatomical Abbreviations 


A-anus; an-anterior nostril; AR- anal fin ray; 
B, - first basibranchial; BO - basioccipital; BR - 
branchiostegal ray; BS-basisphenoid; Cy, - first 
ceratobranchial; CE-centrum; CH - ceratohyal; 
Cl-cleithrum; Co-coracoid; CTP-transverse pro- 
cesses of caudal vertebrae; CX - cartilaginous ex- 
tension of terminal vertebra; D-dentary; DFO - 
dorsal fin origin; DR - dorsal fin ray; E- eyeball; 
E, - first epibranchial; EH - epihyal; EN - epineur- 
al; EO - epiotic; ET - ethmoid portion of premax- 
illoethmovomer; EX - exoccipital; F - frontal; GB - 
gas bladder; GH - glossohyal; GO - gill opening; 
H - heart; H: - first hypobranchial; HA - haemal 
arch; HH-hypohyal; HY-hypural; HYM - hyo- 
mandibular; I - intestine; I, - second infrapharyn- 
gobranchial; IM - intramuscular bone; io - infra- 
orbital pore; IO - interopercle; L/D - relation of 
length to depth; LL- lateral line; LP - lower 
pharyngeal tooth plate; MX - maxilla; N - nasal; 
NA - neural arch; NS - neural spine; OP - opercle; 
OR - orbit; P- parapophysis; PA - parietal; PAS - 
parasphenoid; PD- pneumatic duct; PG - ptery- 
goid; PL- pleural rib; pm - preoperculomandibu- 
lar pore; PO-preopercle; pop - preopercular 
pore; por-postorbital pore; POR - postorbital; 
PR - pectoral fin rays; Pt-pterygiophore; PT - 
pterotic; PTS - pterosphenoid; Q-quadrate; S- 
stomach; SA- sagitta; Sc-scapula; SCI - supra- 
cleithrum; so - supraorbital pore; SO - supraocci- 
pital; SOC -supraoccipital crest; SOP - subop- 
ercle; sp-surface sensory papillae; SP - sphen- 
otic; stp-supratemporal pore; tp-temporal pore; 
UH - urohyal; UP - upper pharyngeal tooth plate; 
V - vertebra; VO - vomer. 


Distributional Abbreviations 


EA -eastern Atlantic; EP-eastern Pacific; IP - 
central and Indo-west Pacific; M - Mediterranean; 
WA - western Atlantic. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Institutional Abbreviations 


The following abbreviations are used in reference 
to material examined: 

ANSP - Academy of Natural Sciences of Philadel- 
phia; BPBM - Bernice P. Bishop Museum; CAS - 
California Academy of Sciences; DANA - Carls- 
bergfondets DANA-Ekspeditioner, Marinbiologisk 
Laboratorium Charlottenlund Slot, Denmark; IA - 
Australian Museum at Sydney; LACM - Los An- 
geles County Museum; MCZ- Museum of Com- 
parative Zoology, Harvard University; MNHN - 
Muséum National d’Histoire Naturelle, Paris; 
MSNG - Museo Civico di Storia Naturale, Genoa, 
Italy; MZUSP - Museu de Zoologia, Universidade 
de Sao Paulo, Sao Paulo, Brazil; RU - Rhodes 
University, Grahamstown, South Africa; SIO - 
Scripps Institution of Oceanography; SU - Stan- 
ford University, also listed as SNHM for Stanford 
Natural History Museum, specimens now de- 
posited at the CAS; TABL- Southeast Fisheries 
Center, Miami, Florida; UCLA - Fish Collection, 
Department of Zoology, University of California 
at Los Angeles; UMML- Rosenstiel School of 
Marine and Atmospheric Science of the Univer- 
sity of Miami; USNM - National Museum of Nat- 
ural History, Smithsonian Institution, Washing- 
ton, D.C.; ZMA- Zodlogisch Museum Amster- 
dam. 


Material Examined 


Listed below is the material utilized for osteo- 
logical examination in this study. The specimens 
are grouped by tribes and listed alphabetically 
within each tribe. Following each specific name 
is the museum abbreviation, museum catalogue 
number, number of specimens, range of the total 
length(s) of the specimen(s) involved. Abbrevia- 
tions are: CS, stained and cleared by the Taylor 
(1967) trypsin technique; GA, gill arches re- 
moved, stained and cleared; H, hyoid removed; 
S, skull preparation; X, radiograph. Specimens 
utilized only for vertebral counts are not in- 
cluded in this listing. 


Ophichthidae - Benthenchelyini. Benthenchelys 
cartieri, DANA 3735, 2(105-115mm), CS. 


Ophichthidae - Myrophini. Ahlia egmontis, SIO 
67-87, 1(268), S; SIO 71-266, 1(337), CS. Muraen- 
ichthys chilensis, SIO 65-645, paratype, 1(248), 
CS; SIO 65-655, paratype, 1(276), CS, 1(292), S. 
Muraenichthys gymnopterus, SIO 69-276, 1(129), 
S, 1(144), CS. Muraenichthys gymnotus, SIO 69- 
266, 1(244), CS. Muraenichthys macropterus, SIO 
69-277, 1(181), CS. Myrophis plumbeus, SIO 69- 


15 


371, 1(182), CS. Myrophis uropterus, CAS 13971, 
1(159), CS; BPBM 27209, 1(182), CS. Myrophis 
vafer, SIO 68-242, 1(193), CS, 2(265-325), S. Pseu- 
domyrophis micropinna, SIO 60-72, 1, head and 
trunk only, CS. Pseudomyrophis nimius, ANSP 
110150, 1(350), CS. Schismorhynchus labialis, CAS 
24687, 5(114-137), CS. Schultzidia johnstonensis, 
SIO 69-267, 1(138), CS. 


Ophichthidae - Callechelyini. Aprognathodon 
platyventris, SIO 68-393, paratypes, 2(312-330), 
CS. Callechelys bilinearis, SIO 70-376, 1(260-+), 
CS. Callechelys cliffi, SIO 61-247, 1(218), GA, H, 
X; SIO 65-281, 1(298), GA, H, X. Callechelys eris- 
tigmus, SIO 65-185, paratype, 1(552), GA, H, X; 
SIO 65-354, paratype, 1(431), CS. Callechelys gal- 
apagensis, UCLA 64-40, paratype, 1(767), GA, H, 
X. Callechelys holochromus (holotype of Crypto- 
pterygium holochroma), USNM 154994, 1(801), 
X. Callechelys luteus, SIO 68-497, 1(1038), H, X. 
Callechelys marmoratus, SIO 69-269, 1(286), CS, 
1(340), S. Callechelys melanotaenius, SIO 69-269, 
1(401), CS. Callechelys muraena, TABL Oregon 
2819, 1(235), GA, H, X. Callechelys nebulosus, 
SIO 71-197, 1(283), CS. Callechelys springeri (hol- 
otype of Gordiichthys springeri), USNM 121604, 
1(372), X. Callechelys striatus SIO 71-165, 1(430), 
H, X. Letharchus velifer, holotype, USNM 31458, 
1(396), X. Letharchus rosenblatti, SIO 67-40, para- 
type, 1(248), CS. Leuropharus lasiops, holotype, 
SU 57313, 1(174), GA, X. Paraletharchus opercul- 
aris, UCLA 64-38, 1(435), GA, X. Paraletharchus 
pacificus, SIO 65-321, 2(276-369), CS. 


Ophichthidae - Bascanichthyini. Allips concol- 
or, holotype, CAS 13967, 1(375), GA, X. Bascan- 
ichthys panamensis, SIO 71-98, 1(425), CS; SIO 
71-224, 1(295), CS, 1(510), S. Caralophia loxochila, 
SIO 70-228, 1(445), CS; SIO 70-376, 1(238), CS. 
Dalophis imberbis, SIO 72-290, 1(440), GA, X. 
Ethadophis byrnei, holotype, SIO 67-31, 1(508), 
GA, X. Ethadophis merenda, holotype, SIO 65-47, 
1(530), GA, X. Leptenchelys vermiformis, holo- 
type, USNM 101785, 1(115), X. Phaenomonas 
cooperae, CAS 13964, 2(451-549), CS. Phaenom- 
onas pinnata SIO 65-348, 1(375), CS, 1(375), S. 


Ophichthidae - Sphagebranchini. Apterichtus 
caecus, MSNG 41058, 1(435), X. Apterichtus flavi- 
caudus, SIO 69-364, 1(300), CS. Caecula ptery- 
gera, USNM 206375, 1(232), CS. Cirricaecula 
johnsoni, paratype, USNM 141189, 1(325), CS. 
Ichthyapus ophioneus, SIO 70-376, 1(337), GA, X. 
Ichthyapus selachops, SIO 61-232, 1(400), CS, 
1(400), S; SIO 65-343, 1(234), CS. Ichthyapus vul- 
turis, holotype, ZMA 104.153, 1(240), X; SIO 69- 


16 


366, 1(446), GA, X. Lamnostoma kampeni, SU 
24593, 1(435), GA, X. Lamnostoma orientalis, CAS 
13959, 1(205), CS; CAS 13968, 1(229), CS. Stictor- 
hinus potamius, MZUSP 8959, paratype, 1(289), 
CS. Yirrkala kaupi, SU 26827, 1(345), GA, X. Yir- 
rkala lumbricoides, CAS 13969, 1(346), CS; para- 
type of Y. chaselingi, 1A 16190-601, 1(560), X. 
Yirrkala misolensis, CAS 13965, 1(335), CS. Yir- 
rkala tenuis, SIO 71-165, 1(370), CS. Yirrkala sp., 
BPBM 11858, 1(306), CS. 


Ophichthidae - Ophichthini. Aplatophis chau- 
liodus, UMML 27209, 1(165), CS. Brachysomophis 
sauropsis, SIO 69-267, 1(323), CS; SIO 69-271, 
1(197), CS. Cirrhimuraena macgregori, SIO 68- 
434, 2(291-317), CS. Cirrhimuraena taeniopterus, 
CAS 13962, 1(445), GA, H, X. Elapsopis cyclor- 
hinus, SIO 69-267, 1(268), CS, 1(475), GA. Eche- 
lus myrus, SIO 69-369, 1(449), dissected. Echelus 
pachyrhynchus, SIO 69-370, 1(355), CS. Echiophis 
sp., UMML 29144, 1(298), CS. Evips percinctus, 
holotype, CAS 13966, 1(125.5), GA, X. Leiuranus 
semicinctus, SIO 61-132, 1(195), CS; SIO 69-268, 
1(243), CS; SIO 69-273, 1(344), S. Malvoliophis 
pinguis, |A 3646, 1(470), CS. Myrichthys colub- 
rinus, SIO 69-272, 1(345), CS. Myrichthys macu- 
losus, SIO 68-497, 1(390), CS; SIO 69-272, 1(310), 
CS. Myrichthys xystrurus, SIO 65-335, 1(420), S; 
SIO 65-354, 2(243-268), CS. Myrichthys sp., SIO 
34-371, 1(386), CS. Ophichthus altipinnis, CAS 
14647, 1(915), GA, X. Ophichthus cephalazona, 
SIO 69-279, 2(230-330), CS. Ophichthus erabo, 
CAS 13960, 1(480), GA, H, X. Ophichthus ophis, 
SU 51724, dissected. Ophichthus rutidoderma- 
toides, CAS 28727, 1(330), CS. Ophichthus tri- 
serialis, SIO 61-193, 1(230), CS; SIO 69-252, 
1(800), S, prepared skeleton. Ophichthus zopho- 
chir, SIO 60-304, 1(217), CS; SIO 65-166, 1(310), 
S, 1(340), S. Ophisurus serpens, RU 76-78, 1(325), 
CS. Phyllophichthus xenodontus, SIO 69-273, 
2(270-305), CS. Pisodonophis boro, SIO 69-281, 
1(410), CS. Pisodonophis cancrivorus, SIO 69- 
307, 1(345), CS. Pisodonophis cruentifer, MCZ 
34529, 1(235), CS. Pisodonophis daspilotus, SIO 
72-73, 1(251), CS. Pogonophis fossatus, SIO 61- 
227, 2(232-249), CS. Quassiremus evionthas, UCLA 
64-19, 1(283), GA, X. Quassiremus nothochir, SIO 
65-334, 2(271-342), CS. Scytalichthys miurus, CAS 
13970, 1(235), CS. Xyrias revulsus, holotype, SU 
6476, 1(890), GA, X. 


Comparative Material - Non-ophichthids. The 
following material was stained and cleared with 
two exceptions. These, Gymnothorax mordax and 
Muraenesox coniceps, were examined from pre- 
pared skeletons. 


SERIES 4, V. 41, #1 McCOSKER — EELS 


Anguillidae. Anguilla rostrata, SIO 69-254, 
1(210). 

Muraenidae. Anarchias galapagensis, SIO 65- 
345, 1(130). Echidna nebulosa, SIO 59-8, 1(178). 
Enchelycore bayeri, CAS GVF 1957-18, 1(255). 
Gymnothorax castaneus, SIO 65-291, 1(140). 
Gymnothorax mordax, SIO skeletal collection, 
1(ca. 1 meter). Gymnothorax panamensis, SIO 61- 
239, 1(180). Gymnothorax schismatorhynchus, 
CAS GVF 1958-13, 1(265). Muraena lentiginosa, 
SIO 65-354, 1(165). Uropterygius necturus, SIO 
65-302, 1(175). 

Simenchelyidae. Simenchelys parasiticus, SIO 
68-479, 1(295). 

Derichthyidae. Derichthys serpentinus, SIO 60- 
239, 1(140). 

Serrivomeridae. Serrivomer sector, SIO 63-374, 
1(305). 

Nemichthyidae. Nemichthys scolopaceous, SIO 
65-243, 1(440). 

Heterenchelyidae. Pythonichthys asodes, para- 
type, UMML 23481, 1(290). 

Muraenesocidae. Muraenesox coniceps, SIO 
skeletal collection, a large adult. 

Moringuidae. Moringua ferruginea, SIO 68-531, 
1(305). 

Congridae. Ariosoma_ gilberti, SIO 62-709, 
3(175-190). Conger cinereus, SIO 68-531, 1(210). 
Gorgasia punctata, SIO 62-270, 1(365). Taenio- 
conger sp., SIO 62-42, 1(235). 

Xenocongridae. Chlopsis apterus, LACM 32555, 
2(152-155). Kaupichthys hyoproroides, SIO 67-45, 
2(150-190). 


Statistical Methods 


Comparisons of vertebral means and variances 
were made with a standard ‘t’” test. Two com- 
puter-programmed grouping techniques were 
used to compare inter- and intra-generic rela- 
tionships within the Callechelyini. These were 
modifications of a clustering technique, the 
weighted variable group method (WVGM)_ of 
Sokal and Michener (1958), and of a recurrent 
group analysis (REGROUP) devised by Fager 
(1957). Each will be discussed briefly, although 
the original sources should be referred to_ if 
further information is desired. 

Wormuth’s (1971) modification of WVGM, 
used by him in ommastrephid squid taxonomy, 
was used to identify the interspecific relation- 
ships of 18 of the 21 species of the Callechelyini. 
Cluster analysis is a technique widely used by 
numerical taxonomists, wherein each species (or 
individual) is termed an operational taxonomic 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


unit (OTU). The program, as described by Wor- 
muth (1971), operates in the following manner: 


Initially a matrix of m characters by n OTU’s 
is set up and the values in each row are 
standardized using row means and variances. 
From the standardized matrix (mxn) a 
product moment correlation coefficient is 
calculated. This matrix expresses the rela- 
tionships between all possible pairs of 
OTU’s quantitatively. At this point a cluster- 
ing procedure is employed to extract a 
graphic representation of the information 
contained in the correlation matrix. Any pair 
of OTU’s which has a higher correlation with 
each other than either has for any other 
OTU is put together as a group. An average 
correlation coefficient is computed for each 
group and it is, thenceforth, treated as a 
single OTU. In the WVGM a variable num- 
ber of new groups are formed on each cycle. 
At the end of each cycle, correlation coeffi- 
cients are recomputed based on the previous 
matrix. On any single clustering cycle two 
alternatives are available. One permits only 
groups of two OTU’s to form prior to recom- 
putation of the correlation matrix if their 
incorporation lowers the overall group cor- 
relation value by less than a_ preselected 
amount. As the results of both alternatives 
over a number of trials were very similar, 
the latter option was selected for its shorter 
computation time. The levels at which 
groups are formed are plotted. The graphi- 
cal representation of the results is termed a 
phenogram. 


Data and characteristics used in program WVGM 
for the Callechelyini are presented in tables 8 
and 9. 

Recurrent group analysis was devised by Fager 
(1957) to identify communities of species on the 
basis of their co-occurrence in samples (Fager 
and McGowan, 1963; Fager and Longhurst, 1968) 
and later modified for taxonomic purposes by 
Ebeling and Weed (1963) and by Fager (1969). 
For each species pair, the program calculates an 
index of affinity. Fager and McGowan (1963) 
state that this index does not follow the hyper- 
geometric distribution exactly, and have there- 
fore replaced it with the geometric means of the 
proportion of common characteristics, corrected 
for the number of characteristics recorded for 
the species, such that: 


=u Be 


l= J 
AxB 
where | = index of affinity, ) = number of com- 


mon characteristics, A and B = total characteris- 
tics recorded for species A and species B, and 


17 


where B = A. All characteristics are equally 
weighted. From the matrix of species pairs which 
is generated the largest possible group of species 
is selected. A “breakpoint” is selected such that 
pairs of species in which | is greater than or 
equal to that point are considered to show af- 
finity. For example, the selection of 0.500 as a 
breakpoint would group species which share 
somewhat more than “half” their characteristics. 
In this study, breakpoints of 0.500 and 0.600 
were utilized, the latter appearing to give more 
reasonable groups without involving excessive al- 
ternative and unnatural groups. Once selected, 
the largest group is removed and the procedure 
is continued until all possible groups are formed. 
Where two or more groups of equal size are pos- 
sible, the program selects the one for which the 
sum of species pairs’ affinity indices is largest. 
Intergroup relationship can be calculated using 
the sum of characteristics shared by members of 
each group as a fraction of the total possible 
connections (see fig. 44). Inter-group similarity 
is therefore directly related to this fraction. 

It should be noted that the computer programs 
used are each affected by the amount of, and 
manner in which, data are presented. The sensi- 
tivity of each program increases with increased 
data input. REGROUP is particularly insensitive 
to continuous data (e.g., vertebral and branchio- 
stegal ray numbers, body proportions) and treats 
each data interval equally. WVGM, by contrast, 
takes account of continuous data in calculating a 
correlation coefficient, but is somewhat more 
insensitive to dichotomous and_ trichotomous 
data. The characteristics used in each program 
are identified in tables 8 and 9. 


OSTEOLOGY AND FUNCTIONAL ANATOMY 


In the following section the osteology of Oph- 
ichthus zophochir is described and illustrated in 
detail. The bone complexes are treated sepa- 
rately, each beginning with a description of the 
condition of O. zophochir and followed by a 
discussion of variations and_ specializations 
among other genera within the family. Also in- 
cluded in this section are discussions of the oto- 
liths and of the gas bladder and digestive tract 
conditions of certain ophichthids. 

Ophichthus zophochir was selected as the spe- 
cies with which other ophichthids are compared 
because it possesses the majority of ophichthid 
anatomical characters in a rather generalized 
state. This is not meant to imply that O. zopho- 
chir is the most primitive ophichthid, but rather 


18 


that by being generalized, and not specialized 
by means of extreme anatomical reduction, it 
provides a framework for comparison and discus- 
sion. 


Neurocranium 


The neurocranium of O. zophochir is com- 
pletely ossified and well fused along the cranial 
sutures. The skull is stout, rather elongate, and 
truncate posteriorly. The neurocranium of an 
adult O. zophochir, like that of most anguilli- 
forms, is small in relation to the total length of 
the fish. It occupies 5.5 percent of the TL, yet 
the neurocrania of more elongate ophichthids, 
such as Phaenomonas cooperae, occupy as little 
as 1.3 percent. Various aspects of the neuro- 


SERIES 4, V.41, #1 McCOSKER — EELS 


cranium of O. zophochir are illustrated in figures 
2 and 3. Described below are the elements com- 
posing the neurocranium. 
Premaxilloethmovomer. The premaxillae, eth- 
moid, and vomer are fused into a single com- 
plex (PEV) articulating posterodorsally with the 
frontal and posteroventrally with the parasphe- 
noid, and forming the anterior margin of the 
orbit. The anterior portion of the PEV, the pre- 
maxillae, has been shown to be separate from 
the vomer in other eels, including Anguilla an- 
guilla (Norman, 1926), Derichthys serpentinus 
(Beebe, 1935), and Coloconger scholesi (Chan, 
1967), yet in ophichthids there are no distinct 
sutures separating the elements, and their precise 
limits can only be determined ontogenetically. 
The premaxillary portion is expanded in most 


ke 


: rhs 


Figure 2. Neurocranium of Ophichthus zophochir, SIO 65-166. Upper, dorsal view; 
lower, left lateral view. Scale represents 1 mm. Stippled lines represent cephalic lateralis 
canals. Abbreviations are: BO, basioccipital; BS, basisphenoid; E, ethmoid portion of pre- 
maxilloethmovomer; EO, epiotic; EX, exoccipital; F, frontal; N, nasal; OR, orbit; PA, parietal; 
PAS, parasphenoid; POR, postorbitals; PRO, prootic; PT, pterotic; PTS, pterosphenoid; SO, 


supraoccipital; SP, sphenotic; VO, vomer. 


Figure 3. Neurocranium of Ophichthus zophochir, SIO 65-166. Upper, ventral view; 
lower, posterior view. Scale represents 1 mm. Abbreviations are as in Figure 2. Sagitta (SA) 


is outlined by stippled line. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Figure 4. Neurocranium of Benthenchelys cartieri, DANA Sta. 3735. Scale represents 
1 mm. 


Figure 5. Neurocranium of Myrophis vafer, SIO 68-242. Scale represents 1 mm. 
Figure 6. Neurocranium of Muraenichthys chilensis, SIO 65-655. Scale represents 1 mm. 


Figure 7. Neurocranium of Callechelys marmoratus, SIO 69-629. Scale represents 1 mm. 


a9 


SERIES 4, V.41,#1 McCOSKER — EELS 


Figure 8. Neurocranium of Ichthyapus selachops, SIO 65-232. Scale represents 1 mm. 
Abbreviations are: BS, basisphenoid; OR, orbit. 


Figure 9. Neurocranium of Stictorhinus potamius, MZUSP 8959. Scale represents 1 mm. 
Abbreviations are: BS, basisphenoid; OR, orbit. 


Figure 10. Neurocranium of Bascanichthys panamensis, SIO 71-224. Scale represents 
1mm. 


Figure 11. Neurocranium of Myrichthys xystrurus, SIO 65-335. Scale represents 1 mm. 
Abbreviation PTS is for pterosphenoid. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


ophichthids (extremely so in Benthenchelys, fig. 
4) and is toothed in all genera except Aprognath- 
odon. It narrows posteriorly to become the 
vomer, which is toothed in most ophichthids 
(exceptions are Schultzidia, Phyllophichthus, 
Leiuranus, and Leuropharus). The premaxillary 
and ethmoid dentition, here termed intermaxil- 
lary, is continuous with that of the vomer in O. 
zophochir. A gap separates the intermaxillary and 
vomerine dentition of many ophichthid genera, 
and appears to be a useful character to indicate 
relationship (table 1). The vomerine dentition 
does not continue onto the parasphenoid. The 
ethmoid portion forms the anterior margin of the 
orbit and the medial margin of the nasals. The 
ethmoid is perforated anterolaterally by the lat- 
eral commissure of the first cranial nerve. 

Nasals. The nasals of O. zophochir are paired, 
thin, laminar, and cartilaginous along their ex- 
ternal edges. The anterior portion of the supra- 
orbital cephalic lateralis nerve tract passes 
through the canal along the median edge of the 
nasal. The nasals of the Myrophinae are either 
cartilaginous or absent. Nasal development 
among the Ophichthinae is variable. In certain 
long-jawed ophichthines (including Brachysomo- 
phis, Scytalichthys, and Aplatophis) the nasals are 
either rudimentary or absent. Nasal cartilage ex- 
tends from the anterior edge of the PEV; its de- 
velopment is quite variable, generally consisting 
of two short lateral rods; in certain genera it is 
fused centrally. 

Parasphenoid. The parasphenoid (PAS) is a 
long, narrow, toothless bone, anteriorly overlying 
the vomer and forming the ventral margin of 
orbit. Centrally, it is spread laterally to form the 
anterior floor of the cranium, narrows posteri- 
orly, and splits into two short prongs. It forms 
the ventral margin of the orbit, and borders the 
orbitosphenoid, pterosphenoid, prootic, and 
basioccipital. 

Prootic. The paired prootics (PRO) combine 
with the paired basioccipitals and pterotics to 
form the otic bulla. They are small, nearly rec- 
tangular, and highly perforated with numerous 
Openings for the passage of nerves and blood 
vessels. Through the most conspicuous foramen 
passes the hyomandibular trunk of the facial 
nerve (VII). Smaller foramina exist for the pas- 
sage of the orbital artery and the jugular vein. 
The major axis of the PRO is horizontal. The 
PRO are bordered by the parasphenoid medially, 
the pterosphenoid anteriorly, the sphenotic an- 
terolaterally, the pterotic dorsolaterally, and the 
basioccipital posteriorly. 


2il 


Basioccipital. The median basioccipital (BO) 
is a small, irregularly shaped bone which forms 
the posterior portion of the otic bulla. Its major 
axis, in contrast to the prootics, is vertical. It is 
bordered medially by the parasphenoid, anteri- 
orly by the prootics, and dorsally by the pterotics 
and exoccipitals. 

Pterotic. The paired pterotics (PT) are elongate, 
narrow anteriorly, and broadly flared laterally 
and posteriorly, forming the lateral edge of the 
roof of the cranium. The cephalic lateralis nerve 
tract passes through the PT and opens anteriorly 
in the frontal and posteriorly at the posterior PT 
margin. Anteromedially the PT are bordered by 
the frontal, followed medially by the parietals 
and epiotics, anterolaterally by the pterosphe- 
noid and sphenotic, ventrolaterally by the prootic 
and basioccipital, and posteriorly by the exoc- 
cipitals. Posterior to the sphenotics, the PT forms 
a sheet-like eave extending beyond the body of 
the cranium. 

Exoccipital. The paired exoccipitals (EX) form 
the dorsal and lateral margins of the foramen 
magnum. They are sutured along their dorsal 
midline, and extend posterodorsally and laterally 
as a semicircular sleeve around the foramen 
magnum. Ventrally, they contact the basioccipi- 
tal, and dorsally the supraoccipital. The foramen 
of the tenth cranial nerve opens posteriorly along 
the ventral EX-BO border, with the ninth opening 
lateroventrally from a foramen slightly antero- 
lateral to that of the tenth. 

Supraoccipital. The supraoccipital (SO) is single, 
small, square to subrectangular in shape, and lies 
along the posterodorsal cranial midline. In O. 
zophochir the narrow median crest of the SO 
extends posteriorly as a small point. The condi- 
tion is typical of many ophichthids, although in 
certain generic groups, particularly Callechelys 
and related genera, they are rounded along the 
posterior SO margin. The SO extends anteriorly 
beneath the parietals, and is bordered laterally 
by the epiotics, posteriorly by the exoccipitals, 
and anteriorly by the prootics in most genera. 
In Schultzidia the SO has surfaced and has sepa- 
rated the parietals, extending anteriorly to con- 
tact the frontal. 

Epiotic. The epiotics (EO) are paired, thin, and 
subrectangular. In O. zophochir they are bor- 
dered anteriorly by the parietals, laterally by the 
pterotics, and medially by the supraoccipital. 
Posteriorly their major axis is changed from hori- 
zontal to vertical in forming the dorsolateral mar- 
gin of the posterior cranial face bordering the 
exoccipitals. The EO, like the supraoccipital and 


the pterotics, forms a narrow sheet-like eave 
along their posterior margins. 

Parietal. The paired, thin, subrectangular pari- 
etals (PA) overlay the posterior margin of the 
frontal. They are bordered anteriorly by the 
frontal, laterally by the pterotics, posterolaterally 
by the epiotics, posteriorly by the supraoccipital, 
and fused medially. In certain ophichthids which 
have a prominent supraoccipital crest the median 
ridge development begins along the parietal mid- 
line and gradually increases to its posterior pro- 
jection. Castle (1972: fig. 10) has erred in illus- 
trating the PA of Benthenchelys as contacting the 
sphenotic; in no ophichthid did | find this 
juncture. 

Frontal. The frontal (F) is a single long element 
which, along with the epiotics and_parietals, 
forms the roof of the cranium. Ontogenetically, 
the frontal is presumably formed from the fusion 
of paried lateral elements, but in juveniles and 
adults there is no evident suture. In O. zophochir 
the F is ridged posteriorly along the dorsal mid- 
line. In Aplatophis this ridge is developed as a 
sharp crest. Several nerve tracts pass through the 
F, including the anterior tract of the cephalic 
lateralis nerve and the transverse frontal com- 
missure, which is unique to the Ophichthidae. 
The frontal is deeply split anteriorly by the in- 
sertion of the ethmoid portion of the PEV in 
some genera, and bordered anteroventrally by 
the orbit and orbitosphenoid, ventrolaterally by 
the parasphenoid, laterally by the pterotic, and 
posteriorly by the parietals. The dorsalmost post- 
orbital of certain species of Ophichthus, Echio- 
phis, and Brachysomophis is weakly sutured to 
the frontal at the level of the transverse com- 
missure. 

Basisphenoid (orbitosphenoid of others). The 
basisphenoid (BS) is a small, unpaired median 
bone with two lateral wings which forms the 
posteroventral margin of the orbit. It is bordered 
dorsally by the frontal, posteriorly by the ptero- 
sphenoids, and ventrally is supported by the 
parasphenoid (a myodome is not present). The 
BS in certain genera with elongate and depressed 
neurocrania has become narrow and elongate, 
as can be seen in a comparison of Ophichthus 
(fig. 2), Ichthyapus (fig. 8), and Stictorhinus (fig. 
9); 

Robins (1971: 164-165) has noted that the use 
of the term “orbitosphenoid” in other eel studies 
(including Gosline, 1950, 1951, 1952; Regan, 
1912; Robins and Robins, 1967; Trewavas, 1932; 
and others) actually pertained to the BS. Chaba- 
naud (1936) stated that the teleostean BS is not 


SERIES 4, V.41, #1 McCOSKER — EELS 


homologous with that of higher vertebrates and 
proposed the name ‘‘porpitual’” for that bone in 
teleosts. Springer (1968: 43-44) agreed with 
Chabanaud’s conclusions but conserved the name 
“basisphenoid’’ because of its widespread usage 
in ichthyology, an act with which | fully agree. 

Pterosphenoid (=alisphenoid). The small paired 
pterosphenoids (PTS) form the anterodorsal roof 
of the cranium. In O. zophochir they are con- 
cave, turning evenly from a longitudinal axis 
(along the margin of the frontal) to a nearly 
transverse axis which abuts the anterior margin 
of the sphenotic. They are bordered anteriorly 
by the frontal, dorsally by the pterotics, pos- 
teriorly by the prootic and sphenotic, and medi- 
ally by the parasphenoid. The conspicuous fora- 
men along the PTS-pterotic border is the an- 
terior opening of the trigemino-facialis chamber. 

Otoliths. Ophichthid otoliths, like those of 
most anguilliforms, are small, and hence have 
received little attention either on a descriptive or 
a comparative basis. Studies are limited to those 
of Frost (1926), which included illustrations of the 
sagittae of Myrus vulgaris (=Echelus myrus), 
Ophichthus gomesii, and Pisodonophis boro, 
and the photograph of the sagitta of Myrophis 
lepturus in Kotthaus (1968). The sagitta (largest 
of the three otolith pairs) of twelve ophichthid 
species were compared in this study. The asteris- 
cus and lapillus were too small to be of com- 
parative value. Ophichthid otoliths are ovate and 
biconvex, with a shallow sulcus on the medial 
surface. They are particularly distinctive in having 
a shallow ostial channel which opens anteriorly 
rather than turning dorsally and opening from 
the sulcus, as is typical of the Congridae (cf. Friz- 
zel and Lamber, 1962: fig. A; Frost, 1926: figs. 
10-11, 15). The sagittae of Ophisurus and Oph- 
ichthus (fig. 12) possess a short anterior rostrum 
and are more elongate than those of Ethadophis 
and Myrophis, which are roughly circular. Sagit- 
tae of Myrichthys and Echiophis are intermediate 
in shape. 

Dentition. Dentition has often been used as a 
principal character to define and differentiate 
ophichthid genera. The location and shape of 
teeth was found in this study to be an important 
indication of relationship. The dentition of O. 
zophochir represents the generalized ophichthid 
condition in being multiserial, conical, and on 
all tooth bearing bones (vomer, ethmoid, pre- 
maxilla, maxilla, and dentary). Considerable vari- 
ation exists within the family, including the 
elongate fanglike dentition of Aplatophis, the 
molariform or granular dentition of Myrichthys, 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Figure 12. Otoliths (medial face, left sagitta) of several ophichthid species. Anterior 


end up. Sulcus outline inked in. Scale represents 2 mm. 


A. Ophichthus triserialis 

B. Ophichthus zophochir 

C. Ophisurus serpens 

D. Echiophis intertinctus 

E. Ethadophis merenda (orientation uncertain) 


> Be Ge ae 


Myrichthys xystrurus 
Echelus pachyrhynchus 
Aplatophis chauliodus 
Myrophis vafer 


i) 
W 


SERIES 4, V. 41, #1 McCOSKER — EELS 


-—H 


Figure 13. Head skeleton of Ophichthus zophochir, SIO 60-304. Scale represents 1 mm. 
Refer to Figures 2-3 (neurocranium), 14 (suspensorium), 17 (hyoid), and 19 (pectoral girdle) 


for names of bones. 


Pisodonophis and certain species of Muraenich- 
thys, the minute, nearly villiform dentition of 
Schultzidia, and the smooth toothless vomer 
of Leiuranus, Leuropharus, Phyllophichthus and 
Schultzidia. Characteristics of the dentition of 
ophichthid genera are summarized in table 1. 


Suspensorium and Jaws 


The conditions of the dentition, suspensorium, 
and jaws are directly related to the feeding habits 
of the various genera. In ophichthids, differences 
in feeding habits (e.g., major differences in prey 
items and adaptations of the predators to the 
different habitats) are greater between the spe- 
cies of different genera than between congeners. 
These differences are well evidenced in the form 
of the suspensorium and jaws of various ophich- 
thids. The juxtaposition of the neurocranium, 
suspensorium and jaws, pectoral girdle and 
hyoid apparatus of O. zophochir is illustrated in 
figure 13. 

All elements of the suspensorium and jaws are 
paired. 

Hyomandibular. The hyomandibular (HYM)_ is 
stout and shaped like an inverted right triangle. 
The HYM of strong-jawed piscivorous genera is 
generally strongly ridged for the attachment of 
the massive adductor mandibularis muscle. The 
dorsal surface of the HYM abuts the sphenotic 
and pterotic. A small irregular condyle along the 
antero-dorsal margin of the HYM fits into a shal- 
low socket formed along the sphenotic-pterotic 
suture. The large process on the posterior HYM 


margin adjoins the anterior process of the op- 
ercle. The ventral portion of the HYM contacts 
the quadrate. 

Quadrate. The small, stout quadrate (Q) is tightly 
sutured to the HYM. The vertical ridge along the 
outer face of the HYM is continuous along the 
Q. Ventrally the Q bears a broad rounded con- 
cave process that contacts the articular bone of 
the mandible. 

Articular. The wedge-shaped articular (AR) is 
narrowed anteriorly, and slides into a pocket 
within the dentary. A remnant of the corono- 
meckelian is present along the inner face of the 
AR of O. zophochir; its presence in other genera 
was not systematically determined. Posteriorly, 
a grooved socket in the AR meets the rounded 
socket of the quadrate. 

Dentary. The dentary (D) is the toothed bone 
of the mandible, joined by the articular pos- 
teriorly and adjoining its opposite member at the 
symphysis by a cartilaginous connection. 

Postorbitals. The three postorbitals (POR) of 
most ophichthids are separate, weak ossicles that 
surround the nerve tract connecting the supra- 
orbital and infraorbital pore tracts. The POR of 
O. zophochir however, are specialized by en- 
largement and fusion to form a continuous strut 
bracing the mandible and neurocranium. This 
specialization, also present in other species of 
Ophichthus and Echiophis, is extreme in Brachy- 
somophis (fig. 15). 

The antorbital cartilaginous strut which Gos- 
line (1952) described for Echelus myrus was not 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Yat 4 flea Oh) 


Wy 
T 
N 


Figure 14. Suspensorium and jaws of Ophichthus zophochir, SIO 65-166. Scale repre- 
sents 5 mm. Abbreviations are: AR, articular; D, dentary; HYM, hyomandibular; IO, inter- 
opercle; MX, maxilla; OP, opercle; PG, pterygoid; PO, preopercle; POR, postorbitals; Q, 
quadrate; SOP, subopercle. 

A. Outer face 

B. Inner face 


26 


SERIES 4, V.41, #1 McCOSKER — EELS 


Figure 15. Right postorbital series of Brachysomophis sauropsis. Scale represents 1 mm. 
Abbreviations are: E, eyeball; MX, maxilla; POR, dorsal-most postorbital; io, infraorbital 


pore tract. 


found in other ophichthids studied (see Remarks 
on Echelus). A similar cartilaginous strut has been 
observed in other eels however, including the 
dysommid Atractodenchelys phrix and in the Syn- 
aphorbranchidae (Robins and Robins, 1970: 307). 
They felt “it (probably) represents either the pre- 
frontal or lateral ethmoid or their fusion.” 

Pterygoid (=palatopterygoid). Pterygoid (PG) 
development is variable within the family, al- 
though a clear pattern of relationship was not 
apparent. The PG is reduced to a narrow splint 
in most ophichthids. In O. zophochir it is thin, 
laminar, pointed anteriorly, and blunt posteriorly. 
It is held in place by the dermal layer and con- 
nective tissue, and contacts neither the quadrate 
nor the hyomandibular. The PG of all ophichthids 
tapers anteriorly, and often posteriorly, is largely 
cartilaginous in many species, and does not ap- 
pear to serve any distinct purpose. Ahlia is ex- 
ceptional in differing from the closely related 
species of Myrophis in the shape of its PG and 
in lacking vomerine teeth (fig. 30). A true pala- 
tine is not present in ophichthids, as shown by 
Robins and Robins (1971) in their discussion of 
the “palatopterygoid arcade.” 

Maxilla. The maxilla (MX) of all ophichthids is 
toothed, elongate, and possesses an anterior 
dorsal process which articulates with the PEV. 
Posteriorly, the maxillae of O. zophochir are 
truncate and do not extend beyond the articular. 
The generalized ophichthid condition however, 


is that of a toothless, elongate, ossified or cartil- 
aginous extension of the MX beyond and lateral 
to the articular. The location of maxillary articula- 
tion with the vomer is affected by the elongation 
of the snout and jaw, and is quite variable within 
the family. The condition is certain ophichthid 
genera is illustrated in figures 16 and 30. 

In concluding this section on the neurocran- 
ium and suspensorium and jaws, certain com- 
ments are in order relating to their specializa- 
tions and functional anatomy. Typical of the pis- 
civorous adaptations of species of Ophichthus, 
Brachysomophis, Echiophis and related species 
are the strengthening provided by the cranial 
vault, the elongate pterotics, the broad junction 
of the hyomandibular along the neurocranium, 
and the bracing of the maxillae to the frontal by 
means of the fused postorbitals. An analagous 
condition exists in the muraenid genus Gymno- 
thorax, in which a postorbital strut strengthens 
the jaws and suspensorium (cf. Burton, 1956: 
fig. 6). Other ophichthid genera, particularly 
among the Myrophinae and elongate ophich- 
thines, are adapted to diets of minute inverte- 
brate prey, and have extremely reduced neuro- 
crania, suspensoria, jaws, and dentition. 


Opercular Series 


The opercular series of ophichthids, and of 
anguilliforms in general, is greatly reduced. This 
reduction is apparently related to the increase 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Figure 16. Maxillary-vomer apposition of several ophichthids. Benthenchelys cartieri 
enlarged 20 times, all others 9 times. 

A. Myrichthys xystrurus D. Callechelys marmoratus 

B. Ophichthus zophochir E. Muraenichthys chilensis 

C. Phyllophichthus xenodontus F. Benthenchelys cartieri 


28 


in number and importance of the branchiostegal 
rays as supporting elements for the branchial 
cavity (Greenwood, et al., 1966). Gosline (1959) 
has correlated the reduction of the opercular 
series, posterior displacement of the gill arches 
and pectoral girdle, and the separation of the 
pectoral girdle from the neurocranium with the 
peculiar branchial pump and circulatory mechan- 
ism of anguilliforms. Opercular reduction and 
deossification of cartilage, and the increase in 
the number of branchiostegals of ophichthids 
appear to be greatest in the smaller myrophines 
and elongate bascanichthyins and sphagebranch- 
ins. Within the Ophichthidae, the condition of 
O. zophochir closely approximates the general- 
ized (primitive) state (fig. 14). The opercle (OP), 
the largest of the series, is posterior to the hyo- 
mandibular and above the subopercle (SOP), 
with its ventral margin lying lateral to the dorsal 
margin of the SOP. The interopercle (lO) over- 
lays the SOP and is below the OP. The pre- 
opercle (PO), smallest of the series, overlays the 
(1O) and contains the tract of the preoperculo- 
mandibular nerve. The SOP and IO are the least 
ossified of the series, although the distal margins 
of most members of the opercular series are car- 
tilaginous in most ophichthids. 

Several specializations in each subfamily are 
useful indicators of phylogeny. For example, the 
SOP is produced posteriorly as a projection en- 
closing the ventral and posterior margins of the 
OP in species of Myrophis (fig. 33), Ahlia, Mur- 
aenichthys (Gosline, 1951a: fig. 3), Pseudomyro- 
phis, and Schismorhynchus. This SOP-OP morph- 
ology is typical of other eel families, including 
certain Congridae (Asano, 1962; Rosenblatt, 
1967), Moringuidae (Trewavas, 1932; Smith and 
Castle, 1972), Xenocongridae (Gosline, 1950, 
1951b; Robins and Robins, 1967), and Xenomy- 
stax atrarius (Peden, 1972). The opercular series 
of Callechelys and related genera is reduced and 
has a conspicuously fringed appearance along the 
margin. The opercular series of Stictorhinus, 
Apterichtus, Ichthyapus, and related genera are 
quite reduced, with elements absent in certain 
genera. 


Hyoid Apparatus 


The hyoid apparatus and the associated branch- 
iostegals provide fundamental characters which 
help to unify the subfamilies within the Ophich- 
thidae. In particular, the broad overlap along the 
ventral midline of the branchial basket is herein 
considered a major phylogenetic character of the 


SERIES 4, V.41,#1 McCOSKER — EELS 


family, not evidenced by homology or converg- 
ence in other eel families. The importance of this 
character, which later led to the combining of 
the Neenchelidae, Echelidae (in part), and the 
Ophichthidae, was recognized by Myers and 
Storey (1939), Gosline (1952), Bertin and Aram- 
bourg (1958), BOhlke (1960) and Nelson (1966b). 

The general usage of the term ‘‘epihyal” in the 
ichthvological literature has been incorrect. As 
Goodrich (1930: 405-406) has pointed out, the 
true epihyal is homologous with the hyomandib- 
ular, and the element incorrectly termed the 
epihyal represents the posterior ossified element 
of the ceratohyal. | am in agreement with Good- 
rich’s conclusions, however in view of its wide- 
spread usage in ichthyology, | have herein used 
the term ‘‘epihyal’”’ to represent the posterior of 
the two ceratohyal elements, and the term ‘‘cera- 
tohyal” for the anterior element. 

The following description of the hyoid appara- 
tus is based on that of O. zophochir (fig. 17A). 
The apparatus consists of the unpaired glossohyal 
(GH) and urohyal (UH), and paired upper hypo- 
hyals (HH), ceratohyals (CH), and epihyals (EH). 
The interhyal is absent. The outer posterodorsal 
margin of the EH is connected by cartilage to 
the inner face of the quadrate and provides sup- 
port for the branchial basket. The CH and EH 
are connected by a stout cartilaginous strut af- 
fording little flexibility along the arch; further 
strengthening is provided by the flanking spike- 
like posterior extension of the CH along the 
outer edge of the EH. The GH is grooved along 
the posterodorsal half and extends forward as a 
small cylindrical teat. The GH and paired upper 
HH are interconnected by a weak cartilaginous 
plate. The upper HH is fused along a broad and 
slightly flexible suture to the CH. A ventrolateral 
extension of the CH braces the HH. The UH, 
lying ventrad to the HH, GH and CH, is flattened 
anteriorly and has a short cartilaginous connec- 
nection from its anterior midpoint to the dorso- 
ventral margin of the GH. Posteriorly, the UH 
extends as a slender ossified spike surrounded 
by a membranous sheath and overlays the 
branchiostegal rays. All the branchiostegal rays 
are inserted on the external face of the arch, 6 
on the CH, 1 on the cartilaginous interspace, and 
19 on the EH. The inner 3 and outer 9 rays are 
closely grouped basally and the remainder are 
more widely spaced. Rays 6-16 are expanded 
slightly at their bases but filiform for the re- 
mainder of their length. The rays broadly overlap 
along the ventral midline in a characteristic man- 
ner. The distal 19 rays of the left EH and cartil- 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


= 


Figure 17. Hyoid arch and branchiostegals of Ophichthus zophochir, an ophichthine, 
and Muraenichthys chilensis, a myrophine. Dorsal view. Scale represents 1 mm. Abbrevia- 
tions are: BR, branchiostegal rays; CH, ceratohyal; EH, epihyal; GH, glossohyal; HH, hypo- 
hyal; UH, urohyal. 

A. Ophichthus zophochir 

B. Muraenichthys chilensis (posterior-most branchiostegals not illustrated) 


29 


30 


aginous interspace are overlain by all the rays of 
the right arch, which are in turn overlain by the 
remaining left CH rays. This pattern of the left 
CH rays overlaying the right CH and EH rays 
which overlap the left EH rays is consistent 
within the Ophichthidae. 

The location and number of branchiostegal 
rays among the genera of ophichthids (table 2) 
and the proximity of the branchiostegal rays to 
the hyoid arch differs markedly in the ophichthid 
subfamilies. In the Myrophinae, which appear to 
follow the generalized anguilliform condition, 
the branchiostegal rays are attached to the outer 
face of the EH, often with a single ray on the 
CH (fig. 17B, Muraenichthys). The remainder of 
the branchiostegal rays, which will hereafter be 
referred to as the ‘accessory branchiostegal 
rays’, are unattached and basally lie well behind 
the hyoid arch. In the Myrophinae, these vary 
from as many as 13 pairs in Benthenchelys 
cartieri to 42 pairs in Myrophis vafer. In the 
Ophichthinae, all branchiostegal rays are at- 
tached to the outer face of the hyoid arch( fig. 
17A, Ophichthus), although in some species the 
rays have secondarily become detached. 

The extreme development of this branchio- 
stegal apparatus is obviously a means of strength- 
ening the gill basket. The manner in which sev- 
eral ophichthines, particularly Ophichthus, Echio- 
phis, Brachysomophis, and Aplatophis have all 
the rays attached to the face of the hyoid is 
probably related to a diet of struggling prey, in 
contrast to the myrophine condition of free rays 
and their diet of comparatively weak prey. The 
reduction of the opercular apparatus and the 
posterior displacement of the entire gill arch 
complex necessitates a supplementary skeletal 
framework to prevent the gill basket from col- 
lapsing during the normal burrowing activities of 
ophichthids. The posterior displacement of the 
gill arches among anguilliform families is extreme 
in the Ophichthidae and the Moringuidae (see 
Nelson, 1966a: fig. 58), both of which comprise 
predominantly sand and mud-burrowing forms. 

The accessory branchiostegal apparatus of 
ophichthid and echelid eels was recognized by 
earlier workers, but Parr (1930) was the first to 
describe it and suggest its function. He created 
the term ‘“‘jugostegalia’’ for the accessory skele- 
ton of the gill cover in species of Myrophis. Be- 
cause of their number he did not consider the 
attached rays to be homologous with the bran- 
chiostegals. In that the accessory rays are un- 
differentiable from the branchiostegal rays, espe- 
cially in those ophichthine species in which all 


SERIES 4, V. 41, #1 McCOSKER — EELS 


rays are attached to the hyoid, | prefer not to use 
Parr’s term, jugostegalia. Problems occur in the 
consideration of jugostegalia as a separate en- 
tity, even if one assumes them to be homologous 
with the branchiostegals. For example, in follow- 
ing Parr’s concept of jugostegalia as those rays 
which are free from the hyoid, McAllister (1968: 
85) stated that “Myrichthys (has) 28 (branchio- 
stegal rays) plus 4 jugostegalia...Myrophis 5 
plus 34-36 jugostegalia.”” McAllister’s (p. 80) re- 
tention of the term jugostegalia ‘‘for those sec- 
ondarily multiplied, overlapping and free bran- 
chiostegals found in certain anguilliforms” is in- 
adequate as seen in the above usage. | therefore 
propose that a more flexible terminology be ap- 
plied to those rays, and prefer the term “‘acces- 
sory branchiostegal rays’’. 

The branchiostegal rays also reflect intragen- 
eric and intergeneric similarities through their 
basal thickening and secondary multiplication. 
The outermost rays of all myrophines and several 
ophichthines are proximally broadened (e.g., 
Muraenichthys, fig. 17B, and Neenchelys, Nelson, 
1966b: fig. 2a). This condition is similar to the 
general anguilliform condition as displayed by 
congrids, muraenids, anguillids, and other apodal 
families. In other ophichthines, particularly Ophi- 
chthus, Aplatophis, Myrichthys, and Quassire- 
mus, the branchiostegals are filiform throughout. 
The basal splitting or secondary multiplication of 
the few inner and outermost rays is probably 
secondarily related to the number and spacing of 
the rays along the arch. For example, in species 
of Echiophis and Ophisurus the branchiostegals 
on the CH are paired, and are often fused at 
their bases. No trends were apparent in this con- 
dition within the Ophichthidae, hence it appears 
to be of limited value as a phylogenetic indi- 
cator. 

The urohyal of most ophichthines is produced 
posteriorly as an ossified spike. The UH of all 
myrophines is limited to an ossified basal plate 
with cartilaginous posterior filaments. The few 
ophichthines that lack the ossified spike are 
clearly derived from the generalized condition 
typified by Ophichthus, rather than from the 
somewhat similar myrophine condition. A cur- 
sory survey of other eel families indicates that 
the ophichthine condition is primitive in relation 
to the more specialized myrophine condition. 

A major subfamilial difference is also evi- 
denced in the ceratohyal. Without exception, the 
CH of the Ophichthinae is split into an elongate 
and pointed distal portion and a shorter, truncate 
medial portion which connects, by means of 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Figure 18. Gill arch skeleton of Ophichthus zophochir, SIO 65-166. Dorsal view. Gill 
arches have been cut along the dorsal midline and spread laterally; left upper and lower 
pharyngeal tooth plates are removed to show underlying bones. Stippling indicates cartilage. 
Scale represents 1 mm. Abbreviations are: By, first basibranchial; Ci, first ceratobranchial; 
Ei, first epibranchial; H:, first hypobranchial; I2, second infrapharyngobranchial; LP, lower 
pharyngeal tooth plate; UPs, upper pharyngeal tooth plate. 


32 


cartilage, with the EH (fig. 17A). The myrophine 
CH is not split, but is rather a simple bowed 
bone, distal to, and terminating at about the 
midpoint, of the EH. The myrophine condition 
appears to involve a unique reduction, whereas 
the ophichthine condition is similar to that of 
the Congridae (Asano, 1962; D. Smith, 1971; 
Rosenblatt, 1967) and other eel families. 

The hypohyals of most ophichthines are like 
those of O. zophochir. In certain otherwise dis- 
similar genera, including Schismorhynchus, Apro- 
gnathodon, and Phyllophichthus, the HH are 
absent. Careful examination of the anterior end 
of the CH did not reveal a suture or line of fu- 
sion, so it might be assumed that the HH are 
lost altogether. Observation of an ontogenetic 
series might provide proof of fusion or absence. 


Gill Arches 


Certain elements of the anguilliform gill arch 
skeleton have been shown to be important in- 
dicators of phylogeny (Nelson, 1966a). The 
Ophichthidae differ from all other eel families 
in a combination of gill arch characters, includ- 
ing: a cartilaginous connection between the 
proximal ends of the dorsal part of the first and 
second gill arches (according to Nelson, peculiar 
to the Ophichthidae); first basibranchial either 
ossified or absent, all others cartilaginous, rudi- 
mentary, or absent; hypobranchials 1-2 ossified; 
second infrapharyngobranchial (ls) ossified. If 
one considers the anguillid or congrid gill arch 
conditions, that of numerous ossified elements 
with minor loss or reduction, to be primitive 
among the anguilliforms, then the ophichthids 
are considerably advanced in having several os- 
seus elements replaced with cartilage, and in 
having others reduced or entirely lost. Extending 
this supposition within the Ophichthidae, the 
Ophichthinae, and in particular the Ophichthini, 
are more primitive than the Myrophinae, which 
have lost the fifth ceratobranchial (C;) and have 
reduced or lost certain basibranchials (tables 3-4). 
The ophichthids are also specialized in having 
the gill arch skeleton displaced posteriorly in re- 
lation to the cranium. Among shallow-water eel 
families this condition is exceeded only in the 
Moringuidae (see Nelson, 1966a: fig. 58). Refer- 
ences to gill arches of ophichthid species are 
limited to Popta’s (1904) pioneering study of 
apodal gill arches (which treated Muraenichthys 
gymnopterus, Leiuranus semicinctus, Caecula 
polyophthalmus, Pisodonophis boro, and Myr- 
ichthys colubrinus), Nelson’s (1966a) detailed 


SERIES 4, V.41, #1 McCOSKER — EELS 


study, which included species from 18 ophich- 
thid genera, and Nelson’s (1966b) treatment of 
Neenchelys buitendijki. 

The following description of the gill arch sys- 
tem of Ophichthus zophochir (fig. 18) illustrates 
the presumably primitive condition within the 
Ophichthidae. Terminology of gill arch elements 
follows that of Nelson (1969: 480). The _ basi- 
branchials are single elements, not intercon- 
nected, lying along the ventral midline; all other 
gill arch elements are paired. The first basibran- 
chial is ossified, slender, and connected by cartil- 
age to the first hypobranchials. Basibranchials 2-4 
are cartilaginous and connected to the adjoining 
hypobranchial pair. Hypobranchials 1-2 are ossi- 
fied and stout. Hypobranchials 3-5 are cartilagi- 
nous, with 4 and 5 fused. Ceratobranchials 1-4 
are ossified and subequal. Ceratobranchial 5 is 
reduced to a slender filament which is fused for 
most of its length with the ventral surface of the 
ventral pharyngeal tooth plate. Epibranchials 1-4 
are short, stout, and bear various processes for 
cartilaginous or ligamentous attachment. The first 
infrapharyngobranchial (l:), as in all eels, is ab- 
sent. The second connects to the first epibranch- 
ial by a cartilaginous strap; this condition, as 
mentioned earlier, is peculiar to all ophichthids. 
The third is “T’’ shaped and distally supports the 
third upper pharyngeal tooth plate (UP3). The 
upper pharyngeal plates are separated by a su- 
ture, the third being much smaller than the 
fourth (UPs). Each plate bears along one margin 
a single row of slightly retrorse conical teeth 
which grades to a fine-toothed pavement. /n situ, 
the tooth plates overlie each other, and surround 
the esophageal canal so that the corresponding 
toothed areas of the upper and lower plates are 
aligned. 

The third hypobranchial is cartilaginous in 
nearly all ophichthids. This specialization prob- 
ably functions to increase the flexibility of the 
gill arch skeleton and, in particular, to allow 
further anterior movement of the lower pharyn- 
geal tooth plates. For the same reason the fourth 
hypobranchial is never ossified. The species with 
ossified third hypobranchials, Dalophis imberbis, 
Aprognathodon platyventris, and Elapsopis cyclor- 
hinus, are highly specialized and not closely re- 
lated. Two examples of each of the latter two 
species were examined to rule out the possibility 
of anomalous specimens. Both specimens of 
Elapsopis had a well ossified Hs pair, and 
slender, nearly subequal tooth plates bearing 
conical biserial teeth. The two specimens of 
Aprognathodon were somewhat aberrant in the 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Hs condition. In one, the left Hz was well ossi- 
fied but the right was cartilaginous. The other 
had small discontinuous pieces of bone lying 
within a cartilaginous matrix on both sides. In 
both specimens the short, stout Hi-2 pairs nearly 
met along the ventral midline, and the small 
tooth plates bore several slender pointed teeth. 
In these exceptional species there has probably 
been a return to an ossified Hs condition rather 
than a retention of the primitive ancestral oph- 
ichthid condition, possibly related to a special- 
ized diet which would require less flexibility in 
the gill arch skeleton. 

Microscopic examination and interpretation of 
the gill arch elements is often difficult, particu- 
larly in differentiating the rudimentary and cartil- 
aginous conditions of the basibranchials. Nelson 
(1966a: 393), for example, considered the fourth 
basibranchial condition of Leiuranus semicinctus 
and Machaerenchelys phoenixensis to differ, al- 
though most authors have regarded the latter 
species as a color variant synonymous with the 
former species. The interpretation of the UP3-UPs 
fusion also tends to be subjective and is not 
heavily relied upon. Since anomalies often occur 
in the gill arch skeleton, duplicate specimens 
were examined in this study when suspicious re- 
sults were encountered. 

Nelson (1966a) identified certain ophichthid 
lineages on the basis of the gill arch conditions. 
These comprised (a) those genera with a moder- 
ately well-developed series of basibranchials and 
an ossified fifth ceratobranchial, (b) those with 
C; reduced or cartilaginous, and (c) those with 
very reduced basibranchials and lacking C;, i.e., 
the Myrophinae. Nelson erred in considering 
Echelus myrus to be an echeline (=myrophine) 
but did recognize the generalized condition of 
its basibranchials and C;. My study has shown 
Nelson correct in his other interpretations and 
now places other genera within his general 
framework. 

The functional significance of the modification 
of the gill arches in apodal fishes was discussed 
by Nelson (1966a). The posterior displacement 
of the gill arches, the reduction and modification 
of osseus elements, and the absence of a firm 
interconnection with the cranium  has_ trans- 
formed their function from prey catching (now 
left to the jaws and cranium alone) to one of 
moving large food items through a secondarily 
elongated pharynx. As Nelson (p. 404) has 
pointed out, ‘many of these same modifications 
have occurred independently among syngnathi- 
form fishes and symbranchiform fishes . . ., pos- 


338) 


sibly also as a result of spatial separation of jaws 
and gill arches.” 


Pectoral Girdle 


The pectoral girdle varies considerably within 
the Ophichthidae, grading from a well developed 
to a reduced condition in both subfamilies. The 
primitive condition, represented by Ophichthus 
(fig. 19A) in the Ophichthinae and by Myrophis 
in the Myrophinae (figs. 20E-F), includes the re- 
tention of the cleithrum (Cl), supracleithrum 
(SCI), scapula (=hypercoracoid) (Sc), and cora- 
coid (=hypocoracoid) (Co), and in the Ophich- 
thinae, the actinosts. The postemporal is absent 
and the pectoral girdle is not attached to the 
cranium in anguilliforms (Gosline, 1971). The pec- 
toral fin is best developed in genera such as 
Ophichthus, Echelus, Echiophis, Mystriophis, Ap- 
latophis, Malvoliophis, Cirrhimuraena, Pisodono- 
phis, and Pogonophis in the Ophichthinae, and 
Myrophis and Ahlia in the Myrophinae. The pec- 
toral fin rays are in some cases (Ahlia and Myro- 
phis) multiply-branched. Pectoral rays vary in 
number from three to four weak stubs in Bas- 
canichthys panamensis to 18 rays in Pogonophis 
fossatus. 

The coracoid and scapula generally lie within 
a cartilaginous plate which provides a rigid sup- 
port for pectoral fin movement. When present, 
the pectoral fin base lies in a dorsoventral plane 
and is broad-based in relation to the fin length 
(extremely so in Myrichthys), offering little ro- 
tational movement. Observations of live Echio- 
phis sp. and Ophichthus triserialis indicate that 
those ophichthines use their pectoral fins to as- 
sist the body musculature in making short dart- 
ing movements during feeding. Observations of 
live Bascanichthys panamensis suggest that the 
highly reduced pectoral fin serves little, if any, 
locomotory function. In those forms lacking pec- 
toral fins, the cartilage, as well as the Co and Sc, 
is reduced or absent entirely. The trend in pec- 
toral girdle reduction is related to the burrow- 
ing habit of many ophichthids. The primary func- 
tion of the pectoral girdle as the structural sup- 
port for the pectoral fin has in burrowing forms 
become one of structural support for the poster- 
ior end of the branchial basket. In no species are 
all girdle elements absent, but in Cirricaecula 
(fig. 19N) and in species of Ichthyapus only the Cl 
remains, and in species of Muraenichthys the 
Cl is a thin curved bone which is nearly identical 
in appearance and in function to the last of the 
accessory branchiostegals. The SCI of several 


34 


Figure 19. 


\ 


SERIES 4, V.41, #1 McCOSKER — EELS 


Pectoral girdle of various representative ophichthine genera. All are shown 


in lateral view, right side, except Cirricaecula which is viewed ventrally. Abbreviations are: 


Cl, cleithrum; Co, coracoid; PR, pectoral rays; Pt, pterygiophores; Sc, scapula; SCI, supra- 
cleithrum. 


Sie G)) ott Ta SG) Sa) 


. Ophichthus zophochir 
. Pisodonophis boro 

. Elapsopis cyclorhinus 
. Myrichthys xystrurus 


Quassiremus nothochir 
Aplatophis chauliodus 


. Caralophia loxochila 
. Bascanichthys panamensis 


Phaenomonas pinnata 
Caecula pterygera 


. Lamnostoma orientalis 


Callechelys marmoratus 


. Aprognathodon platyventris 
. Cirricaecula johnsoni 
. Apterichtus flavicaudus 


Yirrkala tenuis 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


rc at 


ID 
Wy, 


Figure 20. Pectoral girdle of various representative myrophine genera. All are shown 
in lateral view, right side. 


. Muraenichthys gymnopterus 
Muraenichthys chilensis 
. Benthenchelys cartieri 
. Ahlia egmontis 
Myrophis vafer 
Myrophis uropterus 


_ Neenchelys buitendijki (from Nelson, 1966b: fig. 2c), pectoral fin rays not illustrated 
. Pseudomyrophis micropinna 


ae (@) art (eal (o) (@Wesh 


36 


species of Muraenichthys is merely a thin sliver 
(e.g., M. chilensis, fig. 20B) and it is lost alto- 
gether in Schismorhynchus. 

A peculiar Co and Sc condition exists in the 
Callechelyini and several of the Bascanichthyini, 
representing a unique ophichthid specialization 
which is not seen in other apodal fishes. It is 
nearly universal among eels for the Sc to be a 
nearly round bone (flattened slightly on the 
posterodorsal edge) lying above the Co (also 
nearly round but slightly flattened on the post- 
eroventral edge), one or both of which are fen- 
estrated. This is also the generalized ophichthine 
and myrophine condition. The Callechelyini 
possess either one or two small rod-shaped 
bones which are connected by cartilage and are 
oriented horizontally in the normal location of 
the Co (see Aprognathodon, fig. 19M and Cal- 
lechelys marmoratus, fig. 19L). Species of Phae- 
nomonas and Ethadophis, and the sphage- 
branchin Lamnostoma orientalis are similar in 
this condition. The homologies of these two rod- 
shaped bones are not entirely clear, but until 
further evidence is discovered, | will consider 
the anterior bone to be homologous with the 
Co and the posterior one to be homologous 
with the Sc. Pectoral girdle reduction in Myrich- 
thys provides an indictation of its generic homo- 
logies, particularly to species of Pisodonophis. 
The Sc in all species of Myrichthys is lost and the 
Co has lost its dorsal curvature (see fig. 19D). 
This condition is also that of Pisodonophis das- 
pilotus, a species clearly more similar to other 
Pisodonophis than to species of Myrichthys in 
other osteological characters. Further similarities 
in all species of both genera include the shape 
of the Cl and SCI. Other Pisodonophis examined 
have retained the Sc and a complete Co, a con- 
dition more like that of Ophichthus. 

The ancestral condition of those genera which 
entirely lack the Co and Sc is not indicated by 
remnants of cartilage or bone or transitional 
species, and is therefore indeterminable. The 
loss of the Co and Sc in Ichthyapus, Apterich- 
tus, Cirricaecula and Quassiremus may have 
been independent, although the first three gen- 
era are related on the basis of other characters. 


Lateralis System 


The apodal lateralis system and associated 
bones have been shown to be useful indicators 
of relationship, particularly within the Congridae 
(Asano, 1962; D. Smith, 1972). Asano showed 
that the number of pores within the cephalic 


SERIES 4, V.41, #1 McCOSKER — EELS 


canal vary within and between genera of Japa- 
nese congrids. The lateralis system within the 
Ophichthidae was found to differ in a character- 
istic manner at the subfamilial and tribal level. 
It is best developed in the Ophichthinae and re- 
duced in the Myrophinae. The ophichthid later- 
alis system has been described and illustrated on 
several occasions, usually on a species by species 
basis and not in a comparative manner. These 
works include: Allis (1903), Ophisurus serpens 
and Echelus myrus (as Myrus vulgaris); Gosline 
(1951a), Ichthyapus vulturis (as Caecula_ platy- 
rhyncha); Hopkirk (1965), Ophichthus zopho- 
chir; Nelson (1966b), Neenchelys buitendijki; 
Blache (1968), Echelus myrus, E. pachyrhynchus 
and Myrophis plumbeus; Blache (1971), Mystrio- 
phis rostellatus, M. crosnieri, and Echiophis in- 
tertinctus; and Blache and Cadenat (1971), My- 
richthys pardalis, Bascanichthys spp., and Cal- 
lechelys spp. 

The ophichthid lateralis system is divisible into 
seven canals: lateral line, supraorbital, infraorbi- 
tal, preoperculomandibular, temporal, supratem- 
poral commissure, and frontal commissure. The 
canals lie either within certain cephalic bones 
(frontal, pterotic, nasal, preopercle, postorbital, 
and dentary) or weakly ossified tubes (ossicles) 
which are broken at short intervals to provide 
flexibility. The Ophichthidae are distinctive in 
having the right and left sides of the cephalic 
lateralis system connected through the frontal 
and temporal canals. 

The following description of the lateralis sys- 
tem of Ophichthus zophochir, a species which 
shows little reduction and minor specialization, 
illustrates the general ophichthine condition (fig. 
21). The supraorbital canal connects with the 
infraorbital and temporal series posteriorly, and 
with its opposite member dorsally across the 
transverse frontal commissure. A single median 
supraorbital pore is present. Three supraorbital 
pores are associated with the nasal, the central 
pore lying within that bone. The anteriormost 
ethmoidal pore is connected to the supraorbital 
canal by a short cartilaginous connection. There 
are six infraorbital pores. Four lie horizontally 
beneath the eye in an ossified canal posterodis- 
tally overlying the maxilla, followed by two verti- 
cal pores lying behind the orbit and connected 
to a canal passing through the three postorbital 
bones. The supraorbital canal connects posteri- 
orly with the temporal canal and commissure, 
passing through the frontal and pterotics. A 
single median supratemporal pore is flanked lat- 
erally by a pore on each side. The temporal 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


stp so 


Figure 21. Cephalic lateralis system and associated bones of Ophichthus zophochir, 
SIO 60-309. Stippling indicates cartilaginous canal section. Scale represents 1 mm. Abbrevia- 
tions are: an, anterior nostril; es, ethmoid section of supraorbital pores; FC, transverse 
frontal commissure; io, infraorbital pores; LL’, anteriomost lateral line pore; N, nasal bone; 
pm, preoperculomandibular pores; pn, location of posterior nostril; PO, preopercle; por, 


postorbital pores; POR, postorbital bones; so, supraorbital pore; stp, supratemporal pore. 
A. Right side view 
B. Dorsal view 


37 


38 


canal extends posteriorly to the lateral line canal 
and ventrally to the preoperculomandibular canal. 
The preopercular section connects by an ossified 
tube to the preopercle, with two ventrolateral 
pores passing from the bone. A cartilaginous 
connection with the third preopercular pore 
joins the preopercular and mandibular sections. 
By overlying the quadrate and angular-articular 
juncture this cartilaginous connection provides 
the flexibility necessary during jaw movement. 
The seven pores comprising the mandibular 
series are unevenly spaced along the dentary. 
The lateral line (LL) canal extends posteriorly 
from the preoperculomandibular-temporal canal 
juncture, loops above the branchial basket, and 
continues midlaterally to within 0.2 head length 
of the tail tip. There are approximately 150 LL 
pores, 51 of them before the anal origin. Lateral 
line ossicles are moderately ossified, separated 
at each pore, and open along the distal margin 
(fig. 22F). On the medial face of all LL ossicles 
there is a centrally located opening for the nerve. 
Lateral line ossicle development is reduced pos- 
teriorly and absent before the last few caudal 
pores. 

Suprageneric relationships among ophichthid 
genera are indicated by repeating pore patterns 
in certain cephalic canals, particularly in the pre- 
opercular portion of the preoperculomandibular 
canal and in the temporal canal (table 5). The 
generalized ophichthine condition consists of 
three preopercular pores and a single temporal 
pore on either side of the median supratemporal 
pore (as in Ophichthus zophochir, Fig 21, and 
Ophisurus serpens, Fig. 24B). The third preoper- 
cular pore (pop*) is lost in certain Ophichthini 
and all Callechelyini and Bascanichthyini. Cer- 
tain Sphagebranchini are specialized in having 
a fourth preopercular pore (pop*) and a second 
temporal pore (tp’), as in Ichthyapus selachops 
(fig. 24A). This condition is not uniform through- 
out the Sphagebranchini, and may vary between 
and within populations, as evidenced by isolated 
populations of Ichthyapus vulturis (Randall and 
McCosker, 1975). The Myrophinae lack tp? and 
pop*. The pop® is present in Ahlia, Myrophis, 
Pseudomyrophis, Muraenichthys, and Schismo- 
rhynchus, but is absent in Neenchelys, Schultz- 
idia, and Benthenchelys (see Nelson, 1966a, figs. 
21-25, in which Schismorhynchus was called 
Leptenchelys labialis, and Nelson, 1966b, fig. 1a). 

The lateral line ossicles also indicate relation- 
ship in their degree of ossification, separation 
at each pore, and the pore position along the 
canal. Most pores lie below the midline of the 


SERIES 4, V.41, #1 McCOSKER — EELS 


LL canal, although some genera are specialized 
in having the pores located centrally within the 
canal. Lateral line ossicles are nearly solid struc- 
tures in the Sphagebranchini and Callechelyini, 
less substantial in the Ophichthini, and most re- 
duced in the Myrophinae (figs. 22-23). 

Certain genera of the Ophichthini are special- 
ized in having a well-developed free sensory 
neuromast system along the sides and top of the 
head. The elaborate development of these sense 
organs has been generally overlooked. The 
neuromasts are not randomly scattered across 
the head region, but tend to follow distinct pat- 
terns. Nelson (1972) identified these lines of 
papillae in esocids as “‘pitlines’’, which are ap- 
parently homologous to the free sensory neuro- 
masts described herein. Following his terminol- 
ogy, the neuromast lines of Ophisurus serpens 
(fig. 24), beginning at the snout tip, include 
paired subnasal, antorbital, anterior, and cheek 
lines, and a single midline crossing the nape. 
Various degrees of development are also present 
in species of Ophichthus, Echelus, Pisodonophis, 
Quassiremus, Cirrhimuraena, Echiophis, Mystrio- 
phis, and Aplatophis. The neuromasts appear as 
minute papillae and are often difficult to discern 
due to skin rugosity and a waxy precipitate that 
forms on preserved specimens. 

Neuromast development is probably related to 
the soft bottom habitat occupied by these spe- 
cies. Schwartz and Hasler (1966) suggested that 
LL pore development of the mudminnow Umbra 
limi is reduced and free neuromasts are devel- 
oped in response to its habit of digging into 
soft mud substrate. In doing so, they suggested, 
the pores of the LL canal could become impacted 
with mud and severly impaired. The numerous 
and widely distributed superficial organs, how- 
ever would remain functional. Rosenblatt and 
Rubinoff (1972: 362) inferred a similar adapta- 
tion in a heterenchelyid eel in noting that 
“the absence of lateral-line pores in P. asodes 
indicate that it may be a burrower in soft mud 
or in the semiliquid mud-water interface.” This 
inverse relation between sensory neuromasts and 
LL canal development is further evidenced by the 
ophichthids in that (a) a general correlation ex- 
ists between neuromast development and a mud, 
rather than a sand, substrate occupied by the 
species involved, (b) free neuromast development 
is absent in the Sphagebranchini where cephalic 
pore development is greatest, and (c) conversely, 
the mud-dwelling species of Echelus display ex- 
treme pore reduction and moderate neuromast 
development. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


me iii a 


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SS (ESS Ese aie reenae | aly IE), 
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Figure 22. 


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Lateral line ossicles of representative ophichthines. Distal view, right side 


of mid-trunk region. Scales represent 1 mm. Stippling indicates lateral line canal. 


. Aplatophis chauliodus 

. Quassiremus nothochir 

. Phyllophichthus xenodontus 

. Aprognathodon platyventris 
Myrichthys xystrurus 
Ophichthus zophochir 

. Cirrhimuraena taeniopterus 

. Paraletharchus pacificus 
Callechelys eristigmus 
Pisodonophis cancrivorus 


= 6 os) ol bal ) eyes oe 


K. Phaenomonas pinnata 
Ethadophis byrnei 

. Ichthyapus selachops 

. Yirrkala tenuis 

. Caralophia loxochila 
Bascanichthys panamensis 

. Allips concolor 
Cirricaecula johnsoni 
Lamnostoma orientalis 


iE 
M 
N 
O 
P. 
Q 
R. 
Se 
T. Elapsopis cyclorhinus 


40 SERIES 4, V.41,#1 McCOSKER — EELS 


_ — hi : = 


TS OS TY eR a al era 


© 
A 
ey eS eS Casas 2S eS ee a eee 
: : 
| a 
=] = 
ae = — <- US o = o.2 EOS OSSS oP PSS ee oe 50° 
Ss 
inched oe be Co ft eS eS Ea aS Sue rs 09s ed De ee ee a 
= 4 —— ——= 
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trey © Hs Ra 


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Figure 23. Lateral line ossicles of representative myrophines. Distal view, right side of 


mid-trunk region. Scales indicate 1 mm. Stippling indicates lateral line canal. 


A. Pseudomyrophis nimius E. Pseudomyrophis micropinna 
B. Myrophis vafer F. Ahlia egmontis 

C. Benthenchelys cartieri G. Schismorhynchus labialis 

D. Muraenichthys chilensis H. Schultzidia johnstonensis 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Figure 24. Cephalic pore and surface sensory papillae development in two ophichthids. 
Abbreviations are: an, anterior nostril; pop!, first preopercular pore; sp, surface sensory 
papillae (free neuromasts); stp, supratemporal pore; tp', first temporal pore. 

A. Ichthyapus selachops, SIO 65-290. Dotted lines represent contours, not surface 

sensory papillae. 

B. Ophisurus serpens, unnumbered Rhodes University specimen, adapted from Allis 

(1903), left lateral view. 
C. Ophisurus serpens, dorsal view. 


41 


42 


Papillae also occur on the snout and tail tip of 
several ophichthids (cf. Rosenblatt and McCosker, 
1970). Their function has neither been examined 
nor proposed, but might be inferred from ob- 
servations and analogous structures on other 
fishes. Tail tip papillae are generally small and 
may function as contact sensory devices in rela- 
tion to the tail-first digging behavior of most 
observed ophichthids. Certain of the snout pa- 
pillae are often larger (e.g., in Leuropharus lasi- 
ops and Evips percinctus) and may serve a gusta- 
tory as well as a contact sensory function. These 
papillae are best developed on snouts of several 
of the small-eyed permanent burrowing species 
(including species of Phaenomonas,  Bascan- 
ichthys, and Ichthyapus). A gustatory function for 
similar papillae on the snout of a heterenchelid 
eel might also be inferred from Rosenblatt and 
Rubinoff’s (1972: 362) description of Python- 
ichthys asodes. They observed a reduction in the 
olfactory epithelium and the development of 
papillae on the jaws of this small-eyed species, 
and suggested this was related to a fossorial 
habit. Most species of ophichthids have not re- 
duced their olfactory epithelium, but probably 
encounter environmental problems similar to 
those faced by Pythonichthys in their modes of 
feeding. 


Axial Skeleton 


Regan (1912) considered the axial skeleton to 
be of major importance in separating eel famil- 
ies. He separated the Echelidae (considered by 
Regan to include Echelus, Ahlia, Myrophis, Par- 
amyrus, Chilorhinus, Muraenichthys, and Eomy- 
rust) and the Ophichthidae from the Congridae 
on the basis of the formers’ vestigial neural 
spines. He further separated the Echelidae from 
the Ophichthidae on the basis of the weaker ribs 
of the latter family. Gosline (1951a: 302-303) 
clarified Regan’s statements in his discussion of 
the ophichthid axial skeleton. Difficulties in the 
preparation and dissection of the anterior verte- 
brae have precluded their usage in this study in 
a systematically comparative manner. 


The following description is based on the axial 
skeleton of Ophichthus zophochir (figs. 25-26). 
The first vertebral (V) centrum (CE) is reduced 
and not fused to the skull (fig. 25A). Its neural 
arch (NA) extends posteriorly over the second 
V. A lateral flange on the CE is present on the 
second and following trunk vertebrae. The NA 
of V 1-5 are smooth. Along the midline of the 
NA of V 1-12 is a single longitudinal crest which 


SERIES 4, V.41,#1 McCOSKER — EELS 


is split at its posterior margin to form two short 
ridges (fig. 26A). Neural spines (NS) are undevel- 
oped on the trunk vertebrae but become weakly 
developed points posterior to the 6th or 7th 
caudal vertebra. Epineurals (EN), epipleurals (EP), 
and pleural ribs (PL) extend posteriorly from the 
NA and parapophyses (P), and are approximately 
5-7 V in length. The EN and EP of most ophich- 
thids begin at the posterior margin of the neuro- 
cranium. The P of V 1-12 are posteriorly directed 
and increase gradually in length. At approxi- 
mately V 13 the P are symmetrical and shaped 
like normal isosceles triangles. Foramina exist on 
each P, slightly posterior to mid-centrum (fig. 26B). 
The P of V 13-45 (approximate) are homogenous 
in size and shape; the lateral processes of the 
following 5-6 V are reduced. The first caudal ver- 
tebra (at which point the haemal arch begins to 
form) differs markedly in having its P split, the 
upper portion directed laterally to become the 
first caudal transverse process (CTP) and the 
lower directed downward to become the haemal 
arch (figs. 26C-D). The CTP are sharp lateral pro- 
jections, incised at their midlines to the centrum, 
and continuing nearly to the caudal tip. The 
haemal arch closure occurs at approximately the 
10th caudal V. The closure however, is incom- 
plete, and consists of the joining of the posterior 
ends of the haemal spines. Intramuscular (IM) 
bones replace the neural and pleural ribs in the 
caudal region. 

Differences in neural arch shape and sculptur- 
ing are evident in comparing the anteriormost 
five vertebrae of species of the type genus of 
each ophichthid tribe (fig. 25). The NA of V 1-5 
of members of the Callechelyini can be distin- 
guished, at the tribal level, on that basis alone. 
Other characters from the axial skeleton were 
found to be useful indicators of relationship. The 
parapophyses of certain sphagebranchin genera, 
for example, were found to possess a marginal 
process which was lacking in related genera (fig. 
33). Also, the CTP are lacking in most myrophins, 
yet in Muraenichthys and related genera the an- 
terior half of the column is similar to the ophich- 
thine column. Finally, the characteristics of the 
pleural ribs of Ahlia and Myrophis were found 
to differ from that of all other ophichthids in that 
they are limited to the anterior 15-20 vertebrae 
(see Remarks concerning Ahlia and Myrophis). 

Vertebral numbers have been shown to be 
useful characters for the separation of species 
and populations of apodal fishes. Their applica- 
tion to the separation of genera is somewhat 
difficult because of the high degree of overlap 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Figure 25. Anteriormost five vertebrae of representative species from the tribes of 
ophichthids. All are shown in left lateral view. Scale represents 1 mm. Abbreviations are: 
Ce, centrum; EN, epineural; NA, neural arch; P, parapophysis; PL, pleural rib. 


A. Ophichthus zophochir D. Callechelys marmoratus 
B. Stictorhinus potamius E. Myrophis vafer 
C. Bascanichthys panamensis F. Benthenchelys cartieri 


44 SERIES 4, V. 41, #1 McCOSKER — EELS 


CTP 


Figure 26. Trunk and caudal vertebrae of Ophichthus zophochir. Arrows point anteri- 
orly. Abbreviations are: Ce, centrum; CTP, transverse processes of caudal vertebrae; EN, 
epineural; IM, intramuscular bone; NA, neural arch; NS, neural spine; P, parapophysis; PL, 
pleural rib. 
A. 
B. 


Cc 


Anterior view of 14th vertebra. Ribs appear foreshortened due to viewing aspect. 
Ventral view of 14th-16th vertebrae. 


Dorsal view of last precaudal (51st) and anterior five caudal vertebrae (52nd-56th). 
Ribs and IM bones not illustrated. 


Ventral view of vertebrae illustrated in C. Ribs and IM bones not illustrated. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


between genera, and a single mean value cannot 
be applied to a genus. Trends, however, are 
present within genera, and are probably relatable 
to the mode of life and associated anatomical 
specializations of the species involved. The spe- 
cies of Phaenomonas, Allips, and Bascanichthys, 
for example, tend to have increased vertebral 
numbers, primarily in the trunk region. An exam- 
ination of radiographs and gut contents of speci- 
mens of those genera disclosed the presence of 
copious sand and gravel particles in the gut and 
intestine, and the absence of any recognizable 
macroscopic animal material. On that basis as 
well as my observations of live specimens, | sug- 
gest that these eels indiscriminately eat their 
way through the substrate, digesting any utiliz- 
able organic material they encounter. In many 
ophichthids the length of the gut is increased by 
a loop that extends into the tail portion. The gut 
of species of Phaenomonas and Bascanichthys is 
straight, presumably to prevent blockage by sand 
particles passing through the lower tract. The in- 
creased trunk length, which is typical of these 
genera, perhaps reflects this problem, and may 
have been necessary to achieve this feeding 
mode. Certain callechelyins exhibit a similar in- 
crease in trunk vertebrae and a comparable life 
style. 

Vertebral number is also the basis of correlat- 
ing the pelagic leptocephalus with the trans- 
formed adult stage of each eel species. Included 
in table 6 are the vertebral numbers of eel spe- 
cies examined and radiographed in the course 
of this study as well as several literature records 
which are assumed to be correct in species 
identification. The literature concerning numbers 
of eel vertebrae has not been exhaustively 
searched in that errors may inadvertently have 
been introduced through improper identification. 


Caudal Skeleton 


Differences in the myrophine and ophichthine 
caudal fins seemed important enough to most 
earlier authors to recognize the lineages as dis- 
tinct families. The Ophichthidae of nineteenth 
century authors was indeed a unique and unified 
assemblage; primarily due to the conspicuously 
pointed tail tip. Important members, most not- 
ably the species of Echelus, were erroneously ex- 
cluded from the Ophichthidae because they 
possessed a weakly developed caudal fin. Gos- 
line (1951a: 303) noted the similarity in the oph- 
ichthine and myrophine caudal skeletons, but 
felt that the continuous median fin condition 


45 


merited subfamilial separation. His findings, to 
my knowledge, have not been questioned by 
subsequent authors. He stated that: 


osteologically, the difference between the 
tails of Muraenichthys and Cirrhimuraena is 
less than that between those of Cirrhimur- 
aena and Caecula platyrhyncha. Rudiment- 
ary rays are present around the tails of both 
Muraenichthys and Cirrhimuraena; they are 
embedded in flesh in Cirrhimuraena (as also 
in Myrichthys). In Caecula platyrhyncha, on 
the other hand, there are no rudimentary 
rays either around the tip of the tail or else- 
where. It is obvious from this discussion... 
that a separate family cannot be maintained 
for Muraenichthys on the basis of tail struc- 
ture. 


The findings of this study are in agreement with 
Gosline’s. Difficulties in the dissection and prep- 
aration of the caudal skeleton has precluded its 
wide usage in this study. Careful examination of 
certain species however (including Echelus my- 
rus, E. pachyrhynchus, Leptenchelys vermiformis, 
and Bascanichthys tenuis), has clarified their 
position within the family. 

The homologies of ossified elements within 
the apodal caudal skeleton are difficult to de- 
termine, and especially so in the case of the 
sharp-tailed ophichthins and sphagebranchins 
that have undergone major modification as an 
adaptation to rapid burrowing. The caudal tip of 
Ophichthus zophochir (fig. 27) is pointed, hard, 
and without visible caudal rays. The median fins 
submerge shortly (approximately one eye dia- 
meter) before the caudal tip. The underlying 
osteology is complex, as is illustrated in Figure 
27. According to the terminology of Rosenblatt 
(1967), which was adapted from Nybelin (1963), 
there is but one weak centrum (CE). Fused to the 
CE is a pointed hypural (HY) and a much reduced 
neural arch (NA). Reduced caudal rays (CR) 
which lack basal elements are imbedded in the 
skin and weakly associated with the HY. The 
caudal skeleton of Myrophis vafer (fig. 28) differs 
somewhat from that of O. zophochir in possess- 
ing a short CE, two elongate HY, and a short 
cartilaginous extension posterior to each HY. The 
caudal rays of Myrophis are split anteriorly to 
receive the hypural plate. This is similar to the 
condition of caudal rays of xenocongrids (Robins 
and Robins, 1967), but appears to differ from the 
relatively unspecialized condition of Anguilla 
(Smith and Castle, 1972: fig. 19a). Blache’s illus- 
tration (1968: figs. 5, 10) of the caudal skeleton 
of Echelus indicates a caudal ray attachment sim- 
ilar to that of Myrophis. Present in most apodal 


46 SERIES 4, V.41, #1 McCOSKER — EELS 


Figure 27. Caudal skeleton of Ophichthus zophochir, SIO 65-166. Scale represents 
1 mm. Abbreviations are: AR, anal ray; B, basal element of pterygiophore; CE, centrum; CR, 
caudal ray; DR, dorsal ray; HA, haemal arch; HY, hypural; IM, intramuscular bone; NA, 
neural arch; R, radial element of pterygiophore. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Figure 28. Caudal skeleton of Myrophis vafer, SIO 68-242. Scale represents 1 mm. Ab- 
breviation CX is for cartilaginous extension of terminal vertebra. 


47 


48 


caudal skeletons is a conspicous foramen be- 
neath the terminal centrum. It is well-developed 
in Myrophis, but reduced or absent in Ophich- 
thus and other ophichthines, resulting from the 
reduction of the lower HY. The haemal aches 
(HA) of the posterior caudal vertebrae of Ophich- 
thus and other ophichthines differs from that of 
Myrophis and other myrophines. The ophich- 
thine condition appears to be that of a simple 
rectangular lateral plate, whereas in the myro- 
phines a wide gap separates the HA into an an- 
terior and a posterior lateral flange. 


The presence of a myrophin-like caudal fin in 
species of Echelus and Leptenchelys requires 
further explanation. As stated above, rudiment- 
ary fin rays are present in the sharp-tailed 
ophichthines. In the discussion of the evolution 
of the Ophichthidae it is hypothesized that the 
elongate bascanichthyins separated early in the 
evolution of the Ophichthinae, and in general, 
they possess a blunt rather than extremely sharp 
pointed tail tip. The tail of Bascanichthys tenuis 
is surrounded by a weak epidermis, which with- 
out careful inspection gives the appearance of a 
rayed caudal fin. The caudal tip of B. tenuis, 
when viewed with transmitted light, was found 
to lack caudal fin rays. The type and only known 
specimen of Leptenchelys vermiformis is similar 
to B. tenuis in possessing loose epidermis at the 
caudal tip, although minute fin rays appear to 
be present. The fin ray development in this juv- 
enile specimen may be anomalous, or may repre- 
sent a redevelopment of the rudimentary fin rays 
characteristic of the ancestral condition. The 
caudal fin of Echelus myrus, in contrast to the 
bascanichthyin fins, has well developed fin rays. 
The caudal skeletons of Ophichthus zophochir 
and £. myrus do not markedly differ other than 
in the development of fin rays. The produced 
rays appear to be a primitive retention of an 
ancestral condition, whereas the hard-pointed 
tail tip of other ophichthines was developed 
early in the evolution of the family. Other primi- 
tive morphological characters of Echelus that 
bear similarities to the generalized ophichthines 
and certain congrids would suggest that Echelus 
is a primitive ophichthid not far from the basal 
ophichthine stock. 


The caudal skeletons of several ophichthids 
have been illustrated by earlier authors. Included 
are: Benthenchelys cartieri (Castle, 1972); Echelus 
myrus, E. pachyrhynchus, and Myrophis plumb- 
eus (Blache, 1968); Mystriophis rostellatus, M. 
crosnieri, and  Echiophis intertinctus (Blache, 


SERIES 4, V. 41, #1 McCOSKER — EELS 


1971); Myrichthys pardalis, Bascanichthys spp., 
and Callechelys spp. (Blache and Cadenat, 1971); 
and Muraenichthys cookei and Cirrhimuraena 
macgregori (Gosline, 1951a). 


Visceral Anatomy 


The digestive tract and gas bladder have been 
shown by Asano (1962) to be useful taxonomic 
characters within the Congridae. This study is 
concerned primarily with osteology and the soft 
anatomy was therefore not examined in a sys- 
tematic manner. 

A cursory examination of a myrophine, Myro- 
phis vafer (SIO 68-286, 240 mm TL), and an 
ophichthine, Ophichthus zophochir (SIO 65-166, 
335 mm TL), disclosed very similar digestive 
tracts and gas bladder morphologies (fig. 29). 
The digestive tract in both species includes a gut 
diverticulum, or stomach (fide D. Smith, 1971), 
which branches off the anterior trunk region and 
extends posteriorly as a blind sac. The intestine, 
in both species examined, is a straight tube 
opening directly into the anus, whereas in other 
ophichthids it appears to extend partially into 
the caudal region and then to loop forward to 
the anus. The gas bladder (GB) connects anter- 
iorly to the intestine through the pneumatic duct 
(PD) at the mid-trunk level. The GB of both 
species is thin walled, surrounded by a thin 
mesentery, and lies alongside the intestine and 
dorsally within the peritoneal cavity. The GB is 
white, shiny and flexible; the digestive tract is 
pale in both species. 

The gas bladder of the above-mentioned spe- 
cies occupies little of the peritoneal cavity, as 
might be expected from their fossorial habits. 
The GB of the pelagic species Benthenchelys car- 
tieri however, is considerably longer than that 
of either of the two fossorial species (Castle, 
1972; fig. 25). Its length is approximately one- 
third of the trunk length, whereas the other two 
are approximately one-fourth to two-ninths. The 
GB of congrid eels is longer than the stomach 
(cf. Asano, 1962; D. Smith, 1971). The GB of 
the three ophichthids, by contrast, is consider- 
ably shorter than the stomach. 


TAXONOMY 


The following section includes an osteological 
definition of the family, a dichotomous key for 
the identification of genera, a diagnosis of the 
subfamilies and tribes, and an osteological and 
external morphological description of each genus 
of the Ophichthidae. 


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49 


Cc 


Figure 29. Comparative anatomy of congrid and ophichthid digestive tract and gas 
bladder. The gas bladder has been separated from the intestine and the mesentery removed 
to improve clarity. The gas bladderx is stippled for identification, not to indicate pigmentation. 
Not drawn to scale. Abbreviations are: A, anus; E, esophagus; GB, gas bladder; H, heart; |, 


intestine; PD, pneumatic duct; S$, stomach. 
A. Ophichthus zophochir, SIO 65-166 


B. Benthenchelys cartieri, after Castle (1972) 
C. Conger myriaster, after Asano (1962) 


The generic key is constructed using both 
osteological and external morphological char- 
acters. An attempt is made to group genera with- 
in the key in a natural manner to indicate rela- 
tionships. 


In this section, redundancy is avoided wher- 
ever possible, however in several instances im- 
portant characters are repeated both in the tribal 
diagnoses and generic descriptions to facilitate 
comparisons. The generic descriptions are based, 
whenever possible, on the type species as well 
as the most morphologically divergent species 
within each genus, in an attempt to include the 
range of variation for each character examined. 
In a few instances the type species of the genus 
was unavailable for osteological study. Those 
genera are identified in the remarks section fol- 
lowing each description. The included nominal 
species of each genus are listed under the head- 


ing “distribution”. Those species known to me 
only from literature records are indicated by an 
asterisk (*). 

Abbreviations of several morphological char- 
acters and conditions are included for the sake 
of brevity. The reader is referred to the listing 
of abbreviations in the Materials and Methods of 
this paper. Also note that Body = Head + Trunk 
when used in body and tail length comparisons. 
The symbol = means ‘‘approximately equal to”. 


Osteological Definition of the Ophichthidae 
From the present study the following osteo- 

logical definition of the Ophichthidae may be 
developed: 

(a) branchiostegal rays numerous and broad- 
ly overlapping along the ventral midline; 

(b) supraorbital canals united by a trans- 
verse commissure through the fused frontals; 

(c) temporal canal present; 


(d) frontals of adults fused for their entire 
length and lacking an obvious suture; 

(e) first epibranchial connected by a con- 
tinuous cartilaginous strap to the second infra- 
pharyngobranchial; 

(f) no more than first basibranchial ossified; 

(g) third hypobranchial usually cartilaginous; 

(h) neural spines rudimentary or absent; 

(i) tongue adnate; 

(j) palatine absent; 

(k) pterygoid well separated from vomer 
and generally free from hyomandibular. 


Analytical Key to the Genera of Ophichthidae 


la. Accessory branchiostegal rays originate be- 
hind ends of epihyal (EH), free rays more 
numerous than attached; caudal fin rays 
conspicuous, confluent with dorsal and 
anal, tail tip flexible; gill openings (GO) 
mid-lateral, a constricted opening .............. 
pe eee ey tee ae IMMWAROVOUIMUIMAG  sacensee teat een 


1b. All branchiostegal rays originate either in 
association with hyoid or before level of 
EH tips; free rays, when present, fewer 
than attached; tail tip a hard or fleshy fin- 
less point; GO mid-lateral to entirely ven- 
tral, un-constricted _...... Ophichthinae........ 9 


2a. 


2b. 


3a. 


3b. 


4a. 


4b. 


SERIES 4, V.41,#1 McCOSKER — EELS 


Neurocranium short, pointed anteriorly, 
broad posteriorly, length/depth = 3; eye 
large, ca. 6 times in head, orbital foramen 
large, its depth ca. 0.5 skull depth; an- 
terior nostril non-tubular; (posterior nostril 
before eye; pectoral fin moderately de- 
veloped) ........ Benthenchelyini .................... 
SRR er ee Oe, NAP ROR Benthenchelys 
Neurocranium more elongate, length/depth 
= 4; eye smaller, 10 or more in head, 
orbital foramen smaller, its depth much 
less than 0.5 skull depth; anterior nostril 


TUMOUR cece cccssnccee My roonitiiip eens 3 
Pleural ribs absent behind 15th-20th trunk 
vertebra; pectoral fin well developed ...... 4 


Pleural ribs present on all trunk vertebrae; 
pectoral fin either absent or moderately 
developed)... eee 5 
Vomerine teeth absent; dorsal fin origin 
(DFO) above or behind anus; maxilla stout, 
not tapering posteriorly, and abutting ptery- 
goid (fig. 30A); hypohyals (HH) absent, 
glossohyal (GH) rudimentary ............ Ahlia 
Vomerine teeth present; DFO anterior to 
mid-trunk region; maxilla thin and tapering 
posteriorly, not closely associated with 
pterygoid (fig. 30B); HH separated from 


Figure 30. Vomer, maxillae, and pterygoid of Ahlia egmontis (A) and Myrophis vafer 
(B). Scale represents 1 mm. Abbreviations are: MX, maxillae; PG, pterygoid. 


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5a. 


5b. 


6a. 


6b. 


ae 


7b. 


8a. 


8b. 


51 


Figure 31. 


ceratohyal by a gap, GH normally devel- 
oped 
Pectoral fin present, coracoid (Co) and 
sometimes scapula (Sc) present; posterior 
nostril lateral; transverse processes of Cau- 
Galieventebraen (GiiR)malbSemts see 6 


Pectoral fin absent, girdle reduced to clei- 
thrum (Cl) and supracleithrum (SCI); pos- 
terior nostril labial; CTP present —.......... 7 


Pectoral fin minute, Sc and actinosts ab- 
Sant 255s ee eeeaee Pseudomyrophis 


Pectoral fin well-developed, Sc, Co, and 
AGCMIMOSUS PORESEMN sce Neenchelys 


Teeth absent on vomer, absent or em- 
bedded on intermaxillary, those on max- 
illa and dentary minute or villiform; supra- 
occipital (SO) extends anteriorly to frontals, 
completely separating parietals; SO crest 
Al SC Il er ie NP RCE tc Schultzidia 
Teeth present on intermaxillary, maxilla, 
dentary, and vomer; SO not extending be- 
VONGmpanletalSaSOmcrest pnesenity asses 8 


A prominent toothed groove on underside 
of snout, bordered by dermal folds, extend- 
ing anteriorly to anterior nostrils; hypohyals 
(HH) fused to ceratohyals (CH); suspen- 
sorium forwardly inclined ....................-------- 


Underside of snout without a prominent 
median toothed groove bordered by der- 
mal folds; HH broadly separated from CH 
by a suture; suspensorium nearly vertical 
Muraenichthys 


9a. 


9b. 


10a. 


10b. 


Wile 


11b. 


12a. 


2b: 


Diagrammatic representation of a species with well developed head pores. 


Neurocranium short, terete, length/depth 
ca. 3 or less; dorsal fin origin (DFO) on 
nape, above supraoccipital (SO); head 
pores reduced pop’, pop‘, and tp? absent 
(fig. 31); hyoid stout, thickened; (gill open- 
ings (GO) inferior, parallel or converging 
forward, isthmus narrower than GO length; 
pectoral fin absent) ...... Callechelyini -..... 10 


Neurocranium longer, length/depth 4 or 
more; DFO, if present, behind nape; head 
pores generally not reduced, may include 
pop’, pop*, tp?; hyoid more slender ...... 14 


Intermaxillary teeth absent; hypohyals (HH) 
absent; third hypobranchial (Hs3) ossified -. 
Aprognathodon 


Intermaxillary teeth present; HH_ broadly 
separated from ceratohyal by a suture; Hs 
Cantilaeiim@ USa tere en ee oe eee lit 


Anterior nostril rim not raised; dorsal fin 
origin (DFO) above epiotics; neurocranium 
slightly depressed, not convex across pari- 
etal-frontal region; four supraorbital pores; 
(anal fin absent; snout not grooved) ........ 
Letharchus 


Anterior nostril tubular; DFO above supra- 


occipital; neurocranium rounded across 
parietals and frontals; three supraorbital 
ONES) 7 at es veceen se tees eee eee ath one eae carareee 72 
Anal fin absent; gill openings (GO) ex- 
panded ventrolaterally, forming broad 
DOGKEtS iis settee ee eee Paraletharchus 
Anal fin present; GO only slightly ex- 


panded, not forming broad pockets ...... 433 


ho 


SERIES 4, V.41, #1 McCOSKER — EELS 


Figure 32. Representation of underside of callechelyin snouts. A. Median groove pre- 
sent (13a in key), as in Callechelys. B. Median groove absent, as in Letharchus. 


Figure 33. Diagrammatic representation of posterior trunk vertebrae (37th), ventral 
view. A. Parapophyses with anterior projection (16a in key), as in Apterichtus. B. Anterior 
projection absent (16b in key), as in Stictorhinus. 


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13a. 


13b. 


14a. 


14b. 


15a. 


15b: 


16a. 


16b. 


lias 


17b. 


18a. 


18b. 


19a. 


Vomer toothed; median groove on under- 
side of snout (fig. 32A); snout and nape 
not heavily papillate -...............-- Callechelys 
Vomerine shaft toothless; no median 
groove on underside of snout; snout and 
nape with numerous papillae ..Leuropharus 
Pectoral fin absent or vestigial; pectoral 
girdle reduced, consisting of a cleithrum 
(Cl), and may include a_ supracleithrum 
(SCI) and reduced scapula (Sc) and cora- 
coid (Co); median fins reduced or absent 
ok 8 2 ae ee ere eo 15 
Pectoral fin present, generally well devel- 
oped; pectoral girdle consists of Cl, SCI 
(except in Scytalichthys), and generally Sc, 
Co, and actinosts; median fins generally 
elevated .............. @phirchithinieeee 29 
Neurocranium depressed and_ elongate, 
length/depth = 4; head pores developed, 
tp? and pop® generally present; gill open- 
ings (GO) entirely ventral (except in Yir- 
nkalal\ ee Sphagebranchini 
Neurocranium deeper and shorter, length/ 
depth = 4; head pores reduced, tp*® and 
pop? absent; GO low lateral, crescentic -... 
Bascanichthyini 
All fins absent; parapophyses of posterior 
trunk vertebrae with an anterior marginal 
projection (fig. 33a); pectoral girdle re- 
duced to a cleithrum and a reduced or ab- 
sent supracleithrum; branchiostegal rays 
few, generally fewer than 20 pairs; second 
basiloranchiall(B>)) absent 22222 all 
Median fins present; anterior margin of 
posterior trunk vertebral parapophyses en- 
tire (fig. 33b); pectoral girdle includes 
cleithrum, supracleithrum, and reduced 
scapula and coracoid; branchiostegal rays 
more numerous, more than 20 pairs; Be 
CaltilalimOUse ee eet * 19 
Upper pharyngeal tooth plates (UP; and 
UP.) fused; cirri present on upper lip ........ 
Cirricaecula 


UP; and UPs separate; upper lip smooth.... 
oe eet irae ee OR a 18 
Posterior nostril opening outside mouth, 
with a flap; anterior nostril tubular; eye 
moderately developed .............. Apterichtus 
Posterior nostril opening inside mouth, 
with or without a flap; anterior nostril flush 
with snout; eye minute .............. Ichthyapus 
Lateral head profile, from above, narrows 
sharply from epiotics to interorbital, then 
extends evenly to a pointed snout; body 


19b. 


20a. 


20b. 


21a. 


21b. 


Daas 


22b. 


Disa 


23b. 


24a. 


24b. 


DE), 


25b. 


53 


stout, its depth less than 30 in its length; 
vomerine teeth enlarged, pointed and re- 
curved Lamnostoma 
Lateral head profile narrows evenly from 
epiotics to snout; body moderately elong- 
ate, its depth more than 40 in its length; 
vomerine teeth conical, not enlarged ....20 
Eye minute, = 5 in snout; anterior nostril 
flush along snout; interopercle (IOP) absent 
4 ens tns 5 Men, SC Re Sa er eae aE Stictorhinus 
Eye larger, = 3 in snout; anterior nostril 
tubular, or with a short but noticeable 
higmcal@© PRE TONES Citi ete oe ee 21 
Neurocranium nearly flat across parietals 
and epiotics; gill openings (GO) with an 
anterolateral duplication forming a pouch; 
accessory branchiostegals loosely attached 
to hyoid, fewer than half associated with 
epihyal (EH); interopercle (IOP) subrec- 
tangular, margin entire -................. Caecula 


Neurocranium raised along dorsal midline, 
not broad and flat across parietals and 
epiotics; gill membrane without a duplica- 
tion; accessory branchiostegals closely as- 
sociated with hyoid, more than half as- 
sociated with EH; IOP rounder, serrated 
along margin Yirrkala 
Tail short, .300-.360 of total length (TL); 
body extremely elongate, its depth ca. 75- 
NGO tina Syw linia eget eee ee eee eee 23 
Tail longer, .395-.530 of TL; body not ex- 
tremely elongate, its depth usually less 
than. OWI ieee ee eee eee 24 
Dorsal fin originating just behind occiput 
and ending less than 2 head lengths behind 
gillvopeningsssanall finmlackinigy: eee 


Bee URE Seo RI shad Lean Aaa Phaenomonas 
Vertical fins low, but extending nearly to 
et atid pp ee oe oe « Hanet ke ge Gordiichthys 
Pectoral fin absent; supraoccipital crest 


(SOC) extends from a parietal ridge, be- 
coming a raised point posteriorly _.......... 25 


Pectoral fin a minute flap in upper gill 
opening corner; SOC nearly rounded, little 
OFMNOMDOStCHOR DO tee ees 28 
Dorsal fin origin (DFO) behind gill open- 
ings (GO); tail longer than body; gill 
arches stout, third hypobranchial (Hs3) ossi- 
fied, fifth ceratobranchial (C;) a slender os- 
SifiedexhOch es ees eee ae et le Dalophis 


DFO above or behind GO; body = tail; 
gill arches reduced, Hs cartilaginous, C; ab- 
sent 


54 


26a. 


26b. 


30b. 


31a. 


31b. 


Bae 


32b. 
33a. 


33b. 


34a. 


34b. 


B5ar 


Anterior nostril not tubular, its rim not 
raised, developed as an opening with lat- 
eral projections into it; underside of snout 
not grooved; intermaxillary teeth incon- 
SDLCUOQUSS ae. pee Fen ee Caralophia 
Anterior nostril tubular; underside of snout 


grooved; intermaxillary teeth conspicuous 
Pt ted Eh ee re Ae 27 
Median fins continuous around caudal, 
caudal fin rays evident ............ Leptenchelys 
Caudal tip blunt, finless -........... Ethadophis 


Dorsal fin origin on head ....Bascanichthys 
Dorsal fin origin more than a head length 


behindeuhiecadi asta aes Pee eA ey Allips 
Fifth ceratobranchial (Cs) absent ............ 30 
Cs present as a slender rod, either ossified 
OT CantilasinOUs essa ee ee eee 34 
Third preopercular pore (pop*) present; 


pectoral fin rudimentary, smaller than eye; 
pectoral girdle reduced to cleithrum and 
SUpraclenthmnUinne ee ee Quassiremus 
pop® absent; pectoral fin well developed, 
longer than eye; scapula and coracoid of 
pectorallcirdlem present, —— 31 
Hypohyals (HH) absent; maxilla with a for- 
ward projection, articulated ca. mid-vomer; 
supraoccipital (SO) rounded, lacking a pos- 
terior projection; urohyal (UH) deeply 
notched anteriorly; anterior nostrils with 
conspicuous leaflike appendages ................ 
Fie Bitte ad OG Soe Se Cree Phyllophichthus 
HH present, separated from ceratohyal by 
a suture; maxilla without anterior projec- 
tions, articulated before mid-vomer; SO 
with a posterior projection; UH not notch- 
ed beyond midpoint of basal plate; an- 
terior nostrils without leaflike appendages 


Pnnenn ann es. Sones $F ONS ol | a AA ey 32 
Jaws subequal; upper pharyngeal tooth 
plates (UP3-UP4) separate _........ Pogonophis 
Lower jaw inferior; UP3-UP4 fused ........ 33 


Third hypobranchial (Hs) ossified; actinosts 
present; vomerine teeth present ..Flapsopis 
Hs cartilaginous; actinosts absent; vomer- 
ines absent, or 1-3 small teeth ....Leiuranus 


Teeth molariform or granular; pectoral fin 
broad-based (fig. 34A) 


Teeth pointed; pectoral fin base restricted, 
opposite upper half of gill openings (fig. 
AID) Wes: be 2 oes amd lee ee 36 
Dorsal fin origin above or behind gill open- 
ings (GO); third preopercular pore (pop?) 
usually present; hypohyals (HH) narrowly 
separated from ceratohyal (CH); supraocci- 


35b. 


36a. 


36b. 


37a. 


37b: 


38a. 


38b. 


39a. 


39b. 


40a. 


40b. 


41a. 


41b. 


42a. 


SERIES 4, V.41,#1 McCOSKER — EELS 


pital (SO) with a posterior projection ........ 
Soa ae see et Re ER Pisodonophis 
DFO well in advance of GO; pop® absent; 
HH broadly separated from CH; SO 
rounded, without a posterior projection... 
Ce ne el a atresia A cman eee Myrichthys 
Eye before middle of upper jaw, preorbital 
portion of neurocranium not extending be- 
yond posterior 2/3, rostral portion of eth- 
moid shorter than orbit; some teeth long 
and_ fanglike 
Eye over middle of upper jaw, preorbital 
portion of neurocranium extends nearly to 
or beyond middle of skull; rostral portion 
of ethmoid about equal in length to orbit; 
teeth inot aan elie eee ne 42 
Lower jaw projects considerably; anterior 
teeth of both jaws long fanglike canines ex- 
tending far outside mouth; frontal crest a 
conspicuous sharp ridge ~.......... Aplatophis 
Lower jaw inferior or jaws nearly subequal; 
anterior teeth in jaws not fangs extending 
beyond snout tip; frontals not forming a 
sharp ridge, neurocranium rounded or flat 
dorsally 
Tail longer than body, compressed posteri- 
orly; pectoral developed, 5 or less in head 
length; third preopercular pore (pop’) pres- 


QNiti «(fee ee 39 
Tail shorter than or nearly equal to body; 
pectoral reduced, 7 or more in head 
leneth a pOp> absentee 40 


Snout short, 7-12 in head length; second- 
ary cephalic papillae absent ........ Echiophis 
Snout longer, 6 or less in head length; 
secondary cephalic papillae well developed 
sh oe gee LY i a Mystriophis 
Postorbitals strongly developed, forming a 
postorbital strut; branchiostegals fewer than 
20; postorbital region with a conspicuous 
transverse depression; lips fringed; colora- 
CON PAU ieee Brachysomophis 


Postorbitals moderately developed, not 
forming a_ strut; branchiostegals 20 or 
more; dorsolateral profile of head even; 
lips entire; body spotted —...................... 41 


Pectoral fin minute, more than 8 in head 
length; body much longer than tail ............ 
wine Es 9 reise Trad Freee te tS ae See Scytalichthys 
Pectoral fin better developed, ca. 7 in head 
length; body and tail nearly subequal -....... 
sik ct eRe ote fs Se Se EE in Xyrias 
Pectoral fin rudimentary, =~ eye; body 
longer than tail; (third preopercular pore 


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Figure 34. Diagrammatic representation of head and pectoral fin of two ophichthins. 
A. Pectorals broad-based (34a in key), as in Myrichthys. B. Pectoral base restricted (34b in 
key), as in Ophichthus. 


55 


56 


42b. 


43a. 


43b. 


44a. 


44b. 


45a. 


45b. 


46a. 


46b. 


SERIES 4, V. 41, #1 


McCOSKER — EELS 


Figure 35. 
in key). 
Fl OSCE LG hehe aoe ee aw eat oe act Selene oe A IR a Evips 
Pectoral fin developed, noticeably longer 
than eye; tail longer than body ............ 43 


Caudal fin present, confluent with dorsal 
and anal; temporal, postorbital, and inter- 
Ofbitallporeswalbsentsse ee Echelus 


Tip of tail a finless point; temporal, post- 
orbital, and interorbital pores present ....44 


Dorsal fin origin (DFO) before gill open- 
ings (GO); third preopercular pore (pop®) 
absent; upper pharyngeal tooth plates (UP3;- 
UP.) fused; pectoral girdle reduced, scap- 
ula (Sc), coracoid (Co), and actinosts ab- 
SC Mitac ete es! sad seks he Malvoliophis 


DFO behind GO, or if before, the upper 
lip is fringed; pop’ usually present; UP3s- 
UP, separate; Sc, Co, and actinosts present 
So UTE ee eI OS ee en ea PE ee Eee 45 


Snout very long, attenuate, ethmoid/neuro- 
cranium = .500; jaws slender and elongate, 
incapable of closing completely in adults .. 
Ophisurus 


Snout moderate or short, ethmoid/neuro- 
cranium < .500 jaws not slender and 
elongate, capable of closing completely..46 


Upper lip not fringed, although a barbel 
may be present; dorsal fin origin (DFO) 
behind gill openings (GO); opercular series 
stout, not weak and serrated along margin; 
aeons wistrlly S4) Ophichthus 


Upper lip fringed (fig. 35); DFO generally 
on head, or above GO; opercular series 
weak, subopercle reduced; actinosts 1-2... 
Cirrhimuraena 


Diagrammatic representation of an ophichthin with a fringed upper lip (46b 


Kaup’s Genera 


Kaup published his generic and specific de- 
scriptions twice in 1856. The earlier treatment, 
“Ubersicht der Aale’’ (1856a), is mentioned by 
John Edward Gray in the preface of the second, 
the Catalogue of Apodal fish in the Collection of 
the British Museum (1856b), published on 30 
December 1856. Gray stated that ‘During the 
printing of the work and the engraving of the 
plates . . . a synopsis of the genera and new 
species has been published by Dr. Kaup, in Ger- 
man, in the Archiv. fur Naturgeschichte, xxii. 41, 
1856." Kaup altered several generic names in 
the latter work (Echiophis became Echiopsis, 
Ophisurapus - Ophisuraphis, and Pisodonophis - 
Pisoodonophis), which has resulted in variant 
spellings of these generic names. Bleeker (1865) 
emended the spelling of several of Kaup’s gen- 
era, but Jordan (1919b) returned to Kaup’s earlier 
work. In the following listings of generic synony- 
mies, the pagination of Kaup’s later work (1856b) 
will follow that of the earlier, set off in paren- 
theses. 


Type species were not designated by Kaup for 
his numerous and short-lived genera. Bleeker 
(1865), as first reviser, synonymized most of 
Kaup’s genera before types were designated. 
Jordan seems to have been the first to designate 
types for Kaup’s invalid genera, and it appears 
that in each case the first species listed by Kaup 
was regarded as the type species (cf. Jordan, 
119 22))r 


Subfamilial and Tribal Diagnoses and 
Generic Descriptions 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Subfamily Myrophinae 


DIAGNOSIS: GO mid-lateral, a constricted open- 
ing; DFO behind mid-trunk; caudal fin rays not 
reduced, externally visible, confluent with dor- 
sal and anal, tail tip flexible; nasals cartilaginous 
or absent; ceratohyal not divided into a short 
median and a long distal portion (fig. 17B); only 
basal plate of urohyal ossified, posterior exten- 
sion cartilaginous; accessory branchiostegal rays 
originate behind tips of epihyal, free rays more 
numerous than attached; branchial skeleton re- 
duced, basibranchials generally limited to first, 
fifth ceratobranchial absent; coloration uniform 
or darkened dorsally. 


Tribe Benthenchelyini 
TYPE GENUS: Benthenchelys Fowler, 1934 


DIAGNOSIS: Body moderately elongate, laterally 
compressed behind head; tail much longer than 
body; anterior nostril not tubular, posterior nos- 
tril lateral, before center of orbit; GO lateral, a 
horizontal ellipse; median fins elevated; pectoral 
fin moderately developed; head pores enlarged, 
a single preopercular pore, frontal commissure 
weakly developed; LL ossicles fragmentary, near- 
ly absent; neurocranium short, rounded (fig. 4); 
otic bulla, nasals, and SOC absent; maxilla broad, 
not produced posteriorly (fig. 16), articulating 
beneath anterior margin of orbit; gill arches 
weakly ossified, B: cartilaginous, Hs ossified, 
UP3;-UP, fused; pectoral girdle moderately de- 
veloped, SCI, Cl, Sc, and Co present; IM bones 
and ribs weakly developed, CTP absent; epi- 
pleurals limited to anterior 14-16 vertebrae; ver- 
tebrae distinctive (fig. 25), neural arches promi- 
nent; caudal more numerous than precaudal ver- 
tebrae; coloration uniform, slightly darker dor- 
sally. Other characters those of the single genus. 


REMARKS: Benthenchelys cartieri was described 
by Fowler (1934) and referred to the Derichthy- 
idae (also Beebe, 1935). Gosline (1952) referred 
it to the Congridae. Subsequently, Castle (1972) 
has recognized it as an ophichthid after a thor- 
ough osteological study. The distinctness of this 
monotypic genus is herein felt to merit tribal 
recognition. The Benthenchelyini appear to be a 
distinctive offshoot from the generalized Myro- 
phini, specialized for a pelagic mode of life. Spe- 
cializations include the large eye, compressed 
body, enlarged median fins, enlarged head pores, 
and slender dentition. These characteristics con- 
verge with those of other pelagic eels, especially 


57 


the genus Derichthys. A myrophin relationship, 
particularly to the generalized Myrophis, is evi- 
denced in the hyoid and branchial arches, gill 
opening, frontal commissure, and disappearance 
of the epipleural ribs. The pelagic life style of 
Benthenchelys (and the associated eye enlarge- 
ment), unique to the Ophichthidae, could have 
evolved from the epipelagic breeding migrations 
of certain myrophines (see Cohen and Dean, 
1970). 


Benthenchelys Fowler 


Benthenchelys Fowler 1934: 267. (Type species; 
B. cartieri Fowler 1934, by original designa- 
tion.) 


DESCRIPTION (supplementing tribal diagnosis): 
snout blunt; jaws nearly subequal; eye large; an- 
terior nostril not tubular, a large anteriad open- 
ing; DFO slightly before vent; jaw and vomer- 
ine teeth conical, recurved, and uniserial, inter- 
maxillary teeth flattened and directed anteriorly, 
separated from those of vomer by a gap; nasal 
cartilage weakly developed; suspensorium an- 
teriorly inclined, jaw angle ca. 95°; maxilla 
broad, not produced posteriorly, articulating be- 
neath anterior margin of orbit; hyoid weak, GH 
elongate, HH separated from CH by a gap, UH 
a subrectangular plate anteriorly, a cartilaginous 
filament posteriorly; branchiostegal rays numer- 
ous, 8 along EH, the last 2 joined basally. 


ETYMOLOGY: From the Greek 3 €2/90f(ben- 
thos), deep, and ‘Ev LEaus (enchelys; either 
masculine or feminine, here to be treated as 
masculine), eel. 


DISTRIBUTION: A single pelagic species (100- 
250 meters) over deep water in the central Indo- 
Pacific. 


Tribe Myrophini 
TYPE GENUS: Myrophis Lutken, 1851 


DIAGNOSIS: Body short to extremely elongate, 
laterally compressed behind head; tail generally 
longer than body; lower jaw included; anterior 
nostril tubular; posterior nostril either lateral or 
labial; GO lateral, a constricted opening; median 
fins low or elevated, DFO behind mid-trunk; 
pectoral fin present or absent; head pores vari- 
ably developed; LL canal weakly ossified; inter- 
maxillary dentition and vomerine, when present, 
continuous; neurocranium not raised along fron- 
tal or parietal midline, SO crest developed in 


58 


some genera; orbit moderately developed; gill 
arches reduced, weakly ossified, Bi often absent, 
Bs.4 absent or rudimentary; pectoral girdle de- 
velopment variable; IM bones and ribs moder- 
ately to weakly developed, transverse processes 
of caudal vertebrae present in some genera; epi- 
pleurals limited to anterior trunk vertebrae in 
some genera; caudal vertebrae more numerous 
than precaudal; coloration uniform, often darker 
dorsally. 


Ahlia Jordan and Davis 


Ahlia Jordan and Davis 1891: 639. (Type species; 
Myrophis egmontis Jordan 1889, by original 
designation.) 


DESCRIPTION: General characters those of My- 
rophis. Differences include: snout sub-conical, 
broad; DFO above or behind anus; vomerine 
teeth absent; maxilla broad, not tapering poster- 
iorly, closely abuts the short and broad pterygoid 
(fig. 30); HH absent, GH rudimentary; gill arches 
reduced, Hs and lz absent. 


ETYMOLOGY: Named for Jonas Nicolas Ahl, au- 
thor of “De Muraena et Ophichtho”, with the 
noun suffix -ia (neuter). 


DISTRIBUTION: A single Caribbean species. 


REMARKS: The controversy regarding generic 
synonymy of Ahlia has never involved a detailed 
osteological study. Those considering it synony- 
mous with Myrophis (Parr, 1930: 8; Hildebrand, 
in Longley and Hildebrand, 1941: 17; Schultz 
and Woods, 1949: 171) did not consider the ab- 
sence of vomerine teeth to represent a generic 
character, but it was assumed by Jordan and 
Davis (1891: 639), Myers and Storey (1939: 158), 
and Wade (1946: 199) that this warranted sep- 
aration. Nelson (1966a: 398) considered Ahlia 
to be distinct on the basis of gill arch characters. 
The generic differences that | have identified are 
clearly related to feeding specialization in A. eg- 
montis, viz., tooth loss, maxillary-pterygoid brac- 
ing, and gill arch reduction, yet the universality 
of these characters among the species of Myro- 
phis suggests that the species of Myrophis form 
a natural group from which Ahlia is a specialized 
offshoot. 

Cohen and Dean (1970) have recorded an in- 
teresting observation of offshore movements and 
a change in eye size accompanying the onset of 
sexual maturity in this species. Their observa- 
tions were made off Honduras, approximately 
145 km from shore. | have made similar observa- 


SERIES 4, V.41, #1 McCOSKER — EELS 


tions within | km from land in the San Blas 
Archipelago, off the Atlantic coast of Panama. 


Muraenichthys Bleeker 


Muraenichthys Bleeker 1853b: 505. (Type spe- 
cies; M. gymnopterus Bleeker 1853, by original 
designation.) 


Scolecenchelys Ogilby 1897: 246. Spelt Scolen- 
chelys by other authors. (Type species; Mur- 
aenichthys australis Macleay 1881, by original 
designation.) 

Myropterura Ogilby 1897: 247. (Type species; 
Myropterura laticaudata Ogilby 1897, by origi- 
nal designation.) 

?Aotea Phillipps 1926: 533. (Type species; Aotea 
acus Phillipps 1926, by monotypy.) 


DESCRIPTION: Body short to moderately elong- 
ate, tail generally longer than body, laterally com- 
pressed posteriorly; snout sub-conical to blunt, 
not deeply grooved on underside; posterior nos- 
tril either along edge of lip beneath a flap or 
opening into mouth; DFO from mid-trunk to 
well behind anus; pectoral fin absent; pop® pres- 
ent; LL ossicles continuous, well developed for a 
myrophin; dentition variable, teeth often multi- 
serial, either conical or blunt, dentition of vomer 
continuous with that of intermaxillary; skull sub- 
truncate posteriorly; SOC present; maxilla elong- 
ate, slender posteriorly (fig. 16); suspensorium 
nearly vertical; opercular series weakly ossified, 
subopercle generally rudimentary, produced pos- 
teriorly in some species (as in Myrophis, fig. 36); 
otic bulla weakly developed; PG short, not brac- 
ing maxillae, reduced and slender in one sub- 
genus; HH separated from CH by a narrow gap; 
gill arches reduced, basibranchials absent, ls os- 
sified and UP;-UPs fusion variable; pectoral 
girdle reduced to a slender Cl and SCI; epipleural 
ribs on all precaudal vertebrae; CTP moderately 
developed. 


ETYMOLOGY: From the Greek se UPA LU a 
an eel, and trots (ichthys; masculine), fish. 


DISTRIBUTION: Nineteen recognized species 
from the tropical, subtropical, and warm temper- 
ate Indo-Pacific Ocean, including a single species 
from the eastern south Pacific. Material exists of 
undescribed Red Sea and western Pacific species. 


REMARKS: Subgeneric lines within Muraenichthys 
were indicated by McCosker (1970) but were 
not designated pending a thorough osteological 
study. My examination and comparison of M. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


gymnopterus, M. chilensis, and M. macropterus 
did not uncover osteological differences compar- 
able to those used to separate other closely re- 
lated ophichthid genera. This result was unex- 
pected in that the external morphology differs 
considerably within the genus, including differ- 
ences in snout form (from blunt to acute), in 
body depth (15-50 times in total length), in the 
number and position of head pores, in dentition, 
and in the character of the posterior nostril. Sub- 
generic lines may be identified in the following 
manner: 


Posterior nostril opens on the outer lip as an 
elongate slit with an anterior flap; a single 
pore between the anterior and_ posterior 
nostrils; jaw teeth usually in bands, inter- 
maxillary teeth in a patch; UP3-UPs fused in 
species examined; snout usually blunt; body 
usually stout, its depth ca. 15-25 in TL ........ 
ee Subgenus Muraenichthys Bleeker 


Posterior nostril opens into mouth, covered 
by an exterior valvular flap; two pores be- 
tween nostrils; jaw teeth uniserial or bi- 
serial, intermaxillary teeth not in a broad 
patch; UP3-UPs separate in species exam- 
ined; snout usually acute; body moderately 
elongate, its depth usually more than 25 in 
see ene. Subgenus Scolecenchelys Ogilby 


The subgenus Muraenichthys includes M. gym- 
nopterus Bleeker (the type species), M. hattae 
Jordan and Snyder, M. schultzei Bleeker, and 
possibly M. macrostomus Bleeker, M. philippin- 
ensis Schultz and Woods, M. sibogae Weber and 
de Beaufort, and M. thompsoni Jordan and Rich- 
ardson. The subgenus Scolecenchelys includes 
M. australis Macleay* (the type species), M. chil- 
ensis McCosker, M. acutirostris Weber and de 
Beaufort, M. cookei Fowler, M. gymnotus Bleeker, 
M. macropterus Bleeker, M. breviceps Gunther, 
and possibly M. iredalei Whitley. Also included 
in Muraenichthys, but not here allocated to a 
subgenus, are Myropterura laticaudata Ogilby, 
Chilorhinus vermiformis Peters, Muraenichthys 
devisi Fowler, M. xorae Smith*, and M. godeffroyi 
Regan. My specimens of M. macropterus dis- 
agreed with Nelson’s (1966a) description in hav- 
ing an unfused UP3-UP4. 

The genera Muraenichthys, Schultzidia, and 
Schismorhynchus display obvious similarities at- 
tributable to a common ancestry. Primitive and 
advanced conditions of certain characters may be 
identified among the approximately 24 species 


59 


involved (many of the species included are 
known to me only from the literature and not 
from specimens). For example, postulated primi- 
tive conditions include the moderately elongate 
body, sub-conical snout, posterior nostril open- 
ing into the mouth, numerous head pores, uni- 
serial or biserial conical teeth, presence of the 
SO crest, posterior development of the sub- 
opercle, separate hypohyals, ossified second in- 
frapharyngobranchial, separate UP3-UPs, and con- 
spicuous cleithrum and supracleithrum. Species 
of the subgenus Scolecenchelys are clearly the 
most primitive, with the species of the subgenus 
Muraenichthys, and Schultzidia and Schismor- 
hynchus as specialized offshoots. The develop- 
ment of transverse processes on the caudal verte- 
brae, shared by these genera, is unique among 
the Myrophinae and without apparent antece- 
dents in more primitive genera such as Myrophis. 

Aotea, type species A. acus, was described by 
Phillipps (1926) on the basis of a partially di- 
gested specimen from New Zealand waters, and 
placed in Muraenichthys by Castle (1967). Whit- 
ley (1968) placed A. acus in the synonymy of 
Muraenichthys breviceps Gunther, yet Phillipps 
(1926: 533-534) characterized A. acus as having 
“fins absent’ and ‘a hard folded portion be- 
neath body posterior to head apparently indi- 
cat(ing) gill-openings...”, both of which would 
exclude Aotea from the subfamily Myrophinae. 
Phillipps’ sketchy description of A. acus does not 
obviously agree with any known _ ophichthine 
genus, but best fits Apterichtus, Ichthyapus, and 
Cirricaecula. Further examination may discover 
that Aotea acus is a species of Apterichtus in 
that the species of the latter two genera are not 
known from even as far south as Australian 
waters. 


Myrophis Lutken 


Myrophis Lutken 1851: 14. (Type species; M. 
punctatus Lutken 1851, by monotypy.) 

Paramyrus Gunther 1870: 51. (Type species; 
Conger cylindroideus Ranzani 1838, by Jordan 
and Davis (1891) as first revisers.) 

Holopterura Cope 1871: 482. (Type species; H. 
plumbea Cope, 1871, by monotypy.) 

Hesperomyrus Myers and Storey 1939: 157. (Type 
species: H. fryi Myers and Storey 1939 = My- 
rophis vafer Jordan and Gilbert, by original 
designation.) 


DESCRIPTION: Body stout to moderately elong- 
ate, laterally compressed throughout; snout sub- 


60 


conical to conical and moderately elongate; eye 
moderate; posterior nostril along edge of lip be- 
neath a flap or opening into mouth; DFO before 
mid-trunk region; pectoral fin moderately devel- 
oped, longer than eye; pop® present; teeth conical, 
uniserial or biserial in jaws and vomer; skull sub- 
truncate posteriorly (fig. 5); SOC present; maxilla 
elongate, slender posteriorly (fig. 30B); sub- 
opercle produced posteriorly as a posteroventral 
border to the opercle (fig. 36); otic bulla weakly 
developed; PG short, not bracing maxilla; Hs 
cartilaginous, UP3-UP, fused in one species; Cl 
and SCI slender, Sc, Co, and an actinost (2) well 
developed; epipleural ribs limited to anterior- 
most 15-20 vertebrae; CTP absent. 


ETYMOLOGY: From the Greek Bu Pos, Myrus, 
and OPCS (ophis; masculine), snake. 


DISTRIBUTION: A circumtropical genus of nine 
nominal species. Included are: Myrophis punc- 
tatus Lutken (WA), M. australis Castelnau (IP)*, 
M. cheni Weng (IP)*, M. lepturus Kotthaus (IP)*, 
M. platyrhynchus Breder (WA)*, M. vafer Jordan 
and Gilbert (EP), Conger uropterus Temminck 
and Schlegel (IP), C. cylindroideus Ranzani (EA)*, 
Holopterura plumbea Cope (EA). Incertae sedis: 
Myrophis frio Jordan and Davis (WA)*. 


REMARKS: Schultz, et al. (1953: 68) erroneously 
included Parabathymyrus Kamohara in the syn- 
onymy of Myrophis. D. Smith (1971) recognized 
it as a valid congrid genus of the subfamily Ba- 
thymyrinae. 


Figure 36. Opercular series of Myrophis vafer, 
SIO 68-242. Right side, distal view. Scale repre- 
sents 1 mm. Abbreviations are: IO, interopercle; 
OP, opercle; PO, preopercle; SOP, subopercle. 


SERIES 4, V. 41, #1 McCOSKER — EELS 


Castle (1963: 16) has discussed the identity of 
the congrid Gnathophis heterognathus (Bleeker) 
Which has been erroneously included in Myro- 
phis by recent authors. 


Neenchelys Bamber 


Neenchelys Bamber 1915: 479. (Type species; 
N. microtretus Bamber 1915, by monotypy.) 


DESCRIPTION: Body moderately elongate, com- 
pressed posteriorly; body shorter than tail; snout 
sub-conical; eye moderate; posterior nostril an 
elongate slit before lower margin of orbit; DFO 
before mid-trunk; pectoral fin moderately de- 
veloped, longer than eye; pop*® absent; teeth 
conical, uniserial except at vomerines and inter- 
maxillary; skull rounded posteriorly; nasal condi- 
tion unknown; SOC absent; maxilla elongate and 
slender posteriorly; subopercle not developed 
posteriorly (fide Nelson 1966b, fig. le); otic bulla 
weakly developed; Bi rudimentary, lz ossified, 
UP;-UP, separate; pectoral girdle developed, SCI, 
Cl, Sc, Co, and an actinost (?) present; epipleu- 
ral rib condition unknown. 


ETYMOLOGY: Presumably from the Greek W€z , 
new, and é UZLEDLS (enchelys; feminine or 
masculine, treated as masculine by Bamber), eel. 


DISTRIBUTION: Two. species, N. microtretus 
Bamber* from the Red Sea, and N. buitendijki 
Weber and de Beaufort* from the Indian Ocean. 


REMARKS: Specimens of Neenchelys were un- 
available for this study. The description is pre- 
pared from Nelson’s (1966b) osteological and 
Mohamed’s (1958) morphological description of 
N. buitendijki. Nelson (1967) noted the presence 
of overlapping branchiostegals in the holotype 
of N. microtretus (apparently the only known 
specimen) but did not compare it with N. buiten- 
dijki. 

Nelson (1966b: 323), in commenting on Wade’s 
(1946) description of Pseudomyrophis micro- 
pinna, stated that ‘‘there is no character signifi- 
cant enough to maintain Pseudomyrophis as a 
genus distinct from Neenchelys. P. nimius, on 
the other hand, seems distinctive enough to be 
placed in a genus of its own.” My osteological 
examination of the species of Pseudomyrophis, 
which | have found to be congeneric, allows 
further comment on this relationship. The two 
genera show certain similarities in morphology 
and habitat (living in mud bottoms in moderately 
deep water) and are more closely related to each 
other than to other genera. Osteological simili- 
larities include the shapes of the neurocrania, 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


pterygoids, maxillae, gill arches, hyoid arches, 
and caudal vertebrae. The characters used in the 
generic key to separate these genera may be ex- 
panded in the following manner: 


DFO in anterior trunk region; snout conical; 
pectoral fin well developed, = snout; pop? 
absent; subopercle small, sub-rectangular; 
(pop? and subopercle condition of N. micro- 
ARERUISMUIMNIKTMOWWIMN)) 22---2020e-002s-c2e 2222-2 Neenchelys 


DFO in posterior trunk region; snout broad, 
tumid; pectoral fin minute, = eye; pop? 
present; subopercle produced posteriorly 
along ventral and posterior margins of op- 
BCS SR ee Pseudomyrophis 


Pseudomyrophis Wade 


Pseudomyrophis Wade 1946: 199. (Type species; 
P. micropinna Wade 1946, by original designa- 
tion.) 


DESCRIPTION: Body moderately to extremely 
elongate, laterally compressed throughout; snout 
broad, tumid; eye small to moderate; posterior 
nostril an elongate slit before lower margin of 
orbit; DFO behind mid-trunk; pectoral fin min- 
ute, smaller than eye; pop’ present; teeth coni- 
cal, uniserial throughout, except at anterior vo- 
mer and intermaxillary; skull rounded _ posteri- 
orly; nasals ossified along canal only, nasal car- 
tilage weakly developed; SOC absent; maxilla 
elongate, slender posteriorly; subopercle similar 
to Myrophis (Fig. 36), produced posteriorly as a 
slender posteroventral border to opercle; otic 
bulla weakly developed; PG short, not produced 
anteriorly; Bi and ls ossified, UP3-UP4 separate; 
pectoral girdle reduced to Cl and SCI (and frac- 
tional Co? in P. micropinna); epipleural ribs on 
all precaudal vertebrae. 


Cha 
ETYMOLOGY: From the Greek W€EUPAYS 
(pseudes), false, -o-, and Myrophis (masculine), 
a genus of ophichthids. 


DISTRIBUTION: Two New World species, P. ni- 
mius Bohlke (Caribbean) and P. micropinna 
Wade (eastern Pacific). 


REMARKS: The species of Pseudomyrophis are 
strikingly different in body depth and head 
length, yet an osteological comparison did not 
uncover differences that are clearly generic. 
Other proportional differences are also related 
to the extreme elongation of P. nimius. The 
mean difference in vertebral number between 
species (ca. 50) is less than that between species 


61 


of Phaenomonas (ca. 70). Both species are ap- 
parently adapted to soft mud bottoms in water 
relatively deep for ophichthids (P. micropinna 
from depths of 45-60 fms, P. nimius to 400 fms). 


Schismorhynchus McCosker 


Schismorhynchus McCosker 1970: 509. (Type 
species; Muraenichthys labialis Seale 1917, by 
original designation.) 


DESCRIPTION: General characteristics those of 
Muraenichthys. Differences include: body mod- 
erately elongate; body shorter than tail; snout 
conical, elongate, with a prominent toothed 
groove on underside; anterior nostril an elong- 
ated tube as long as eye; posterior nostril opens 
into mouth; DFO in posterior trunk region; pop? 
and median interorbital pore absent; teeth coni- 
cal, uniserial; maxilla broad, not becoming 
slender posteriorly; subopercle produced posteri- 
orly as in Myrophis (fig. 36); suspensorium an- 
teriorly inclined; HH fused to CH (or absent?), 
GH rudimentary; gill arches extremely reduced, 
Bi, Hs, and lI» absent, UPs3-UPs fused, lower 
tooth plate elongate (see Nelson, 1966a: figs. 
14-15); pectoral girdle reduced to a slender Cl. 


ETYMOLOGY: From the Greek CLIT UN 
: : O 4 : 
(schisme), cleft, and 7 V zx Of — (latin- 
ized as rhynchus, masculine in accordance with 
item 30(a)(3) of the International Code of Zoo- 

logical Nomenclature), nose. 


DISTRIBUTION: A single species, widespread in 
the central and western Pacific ocean. 


Schultzidia Gosline 


Schultzidia Gosline 1951a: 309. Described as a 
subgenus of Muraenichthys Bleeker. (Type spe- 
cies; Muraenichthys johnstonensis Schultz and 
Woods 1949, by original designation.) 


DESCRIPTION: General characteristics those of 
Muraenichthys. Differences include: body stout 
to moderately elongate, body shorter than tail; 
posterior nostril opens into mouth; DFO behind 
anus; pop® absent; teeth absent on vomer, ab- 
sent or imbedded on intermaxillary, those on 
maxilla and dentary minute or villiform; SOC 
absent, SO extends anteriorly to frontals, com- 
pletely separating parietals; opercular series rudi- 
mentary, subopercle not produced posteriorly; 
PG slender; hyoid rudimentary, HH separated 
from CH by a gap, UH fragmentary, separated 
medially; gill arches extremely reduced, B: and I» 


62 


absent, UPs-UP4 fused, tooth plates elongate and 
vermiculated; CTP weakly developed. 


ETYMOLOGY: Named for Leonard P. Schultz, 
ichthyologist, with the assumed noun suffix -idia 
(feminine). 


DISTRIBUTION: Two central and western Pacific 
species, described as Muraenichthys johnstonen- 
sis Schultz and Woods, and M. retropinnis Seale. 


Subfamily Ophichthinae 


DIAGNOSIS: Gill opening variable, mid-lateral to 
entirely ventral and longitudinal, generally elong- 
ate and unconstricted; DFO variable, from nape 
to behind anus; tail tip generally a hard finless 
point, rudimentary rays visible in certain genera; 
nasals ossified, generally well developed, but re- 
duced or absent in certain genera; CH divided 
into a short median and longer distal portion, the 
median section connecting to the CH by a car- 
tilaginous splint (see fig. 17A); UH generally 
continues posteriorly from basal plate as a 
slender ossified spike; all branchiostegal rays 
originate either in association with hyoid or be- 
fore level of EH tips; gill arch skeleton variably 
developed, first basibranchial always ossified, 
second through fourth generally present in either 
a cartilaginous or rudimentary condition, Cs pres- 
ent in several genera; coloration variable, band- 
ed, barred, striped, spotted, or uniform patterns. 


Tribe Callechelyini 
TYPE GENUS: Callechelys Kaup, 1856 


DIAGNOSIS: Body (head and trunk) and tail 
moderately elongate, laterally compressed; body 
longer than tail; snout acute, rounded at tip; 
lower jaw included; eye small; posterior nostril 
opens into mouth; GO low lateral to entirely 
ventral, converging forward, length much greater 
than isthmus width; dorsal fin originating on 
nape; pectoral fin absent; tail tip a hard finless 
point; head pores reduced, pop*® and tp? absent; 
LL ossicles nearly continuous; teeth conical, jaw 
teeth uniserial, those of vomer separated from 
those of intermaxillary by a gap; skull short, 
sloping posteriorly, its height ca. 3 or less in its 
length (fig. 7); orbit moderately developed; SO 
rounded, without a posterior projection; PG 
slender, elongate, free and tapering posteriorly; 
margin of opercular series irregularly ossified, 
with cartilaginous gaps; suspensorium nearly 
vertical; otic bulla well developed; hyoid stout; 
branchiostegal rays numerous; gill arches re- 


SERIES 4, V.41,#1 McCOSKER — EELS 


duced, C; absent, UP;-UPs separate; pectoral 
girdle reduced to Cl, SCI, and 1 or 2 rod-shaped 
elements; IM bones, ribs, and CTP developed; 
precaudal vertebrae more numerous than caudal; 
coloration variable, either striped, barred, spot- 
ted, mottled, or uniform. 


REMARKS: The Callechelyini constitute the most 
distinct and compact of ophichthid tribes. Avail- 
able material or radiographs of 19 of the 21 spe- 
cies of Callechelyini has allowed an in depth 
study of this tribe. The results of two computer- 
programmed numerical taxonomic evaluations 
of this tribe are presented in the discussion sec- 
tion. Several important morphological and osteo- 
logical characters of the species are listed in 
Table 8. 


Aprognathodon Bohlke 


Aprognathodon Bohlke 1966: 99. (Type species; 
A. platyventris Bohlke 1966, by original desig- 
nation.) 


DESCRIPTION: Anterior nostril tubular; snout 
moderate, rounded at tip; median groove absent 
on underside of snout; intermaxillary teeth ab- 
sent, vomerine teeth present; DFO above SO; 
anal fin present; 3 supraorbital pores; neuro- 
cranium well rounded, highest anterior to front- 
al-parietal suture; hyoid arch very stout, inflex- 
ible along CH-EH suture; HH absent; branchio- 
stegal rays numerous, along arch, often joined 
basally, distal 4-6 rays along EH broadened bas- 
ally; UH a simple cartilaginous projection pos- 
teriorly, basal plate ossified; gill arches stout, Hs 
partially or completely ossified; pectoral girdle 
contains Cl, SCI, and 2 rod-shaped elements; 
body coloration strongly banded longitudinally. 


ETYMOLOGY: From the Greek XH , without, 
rf (pro), forward, VVA eos site 
jaw, and OfavV (odon; masculine), tooth, 

reference to the lack of intermaxillary pe. 


DISTRIBUTION: A single western Atlantic species, 
known from the Bahamas through the Lesser An- 
tilles to Venezuela. 


Callechelys Kaup 


Callechelys Kaup 1856: 51 (28). (Type species; 
C. guichenoti Kaup 1856 = Dalophis marmor- 
ata Bleeker 1853, by monotypy.) 

Cryptopterygium Ginsburg 1951: 482. (Type spe- 
cies; Cryptopterygium holochroma Ginsburg 
1951, by original designation.) 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


DESCRIPTION: Anterior nostril tubular; snout 
short, rounded at tip; a median groove on un- 
derside of snout (Fig. 32A); intermaxillary and 
vomerine teeth present; DFO above SO; anal 
fin present; 3 supraorbital pores; neurocranium 
well rounded, highest anterior to level of frontal- 
parietal suture (fig. 7); hyoid arch stout, moder- 
ately flexible along CH-EH suture; HH separated 
from CH by a narrow gap; branchiostegal rays 
numerous, along arch; distal rays along EH 
broadened basally in some species; UH either a 
simple slender filament posteriorly, or split into 
two divergent rays; Hs cartilaginous; pectoral 
girdle contains Cl, SCI, and either one or two 
rod-shaped elements; coloration variable, either 
uniform, spotted, mottled, or banded. 


ETYMOLOGY: Kaup (1856a, b) did not give the 
derivation of the generic name nor did he desig- 
nate its gender. From his description (1856b: 28), 
“this handsome eel .. .’”, one must assume that 
he intended the generic name to be derived 
from KANNOS (beauty) and € Y LEAVS 
(enchelys, eel) which is feminine, but according 
to Liddell and Scott (1801), was later also mascu- 
line. Bleeker (1865), as first reviser, further con- 
fused matters by recognizing Dalophis marmor- 
ata Bleeker as Callechelys marmoratus and also 
describing Callechelys melanotaenia. To date, 
the gender of Callechelys has not been estab- 
lished, although the most recent revisers (McCos- 
ker and Rosenblatt, 1972) have regarded Calle- 
chelys as masculine. 


DISTRIBUTION: A cosmopolitan genus with 15 
tropical and subtropical species. Nominal species 
include: Callechelys bilinearis Kanazawa (WA), 
C. cliffi Bohlke and Briggs (EP), C. eristigmus 
McCosker and Rosenblatt (EP), C. galapagensis 
McCosker and Rosenblatt (EP), C. /uteus Snyder 
(IP), C. melanotaenia Bleeker (IP), C. muraena 
Jordan and Evermann (WA), C. nebulosus Smith 
(IP), C. perryae Storey (WA, EA), C. striatus Smith 
(IP), Ophichthys bitaeniatus Peters (IP)*, Crypto- 
pterygium holochroma Ginsburg (WA), Caecula 
leucoptera Cadenat (EA)*, Dalophis marmorata 
Bleeker (IP), Gordiichthys springeri Ginsburg 
(WA). 


REMARKS: Numerous authors (Gunther, 1910: 
404; Pellegrin, 1912; Storey, 1939: 63; Smith, 
1957: 83; McCosker and Rosenblatt, 1972: 22) 
have discussed the validity of C. guichenoti, the 
generic type, with the majority supporting its 
synonymy with C. marmoratus. Marie-Louise 


63 


Bauchot of the Paris Museum has kindly furn- 
ished measurements and a radiograph of the 
type specimen (MNHN 2126) of C. guichenoti. 
Its morphometry and osteology (183 vertebrae 
and a single pectoral girdle horizontal element) 
are further evidence of its synonymy with C. 
marmoratus. 

Subgeneric lines within Callechelys were sug- 
gested by McCosker and Rosenblatt (1972). They 
recognized two major groups, one containing 
species with a simple urohyal and a single rod- 
shaped pectoral element (fig. 19L) and another 
with species having the urohyal split posteriorly 
into two slender divergent rays and two rod- 
shaped pectoral elements (as in Aprognathodon, 
fig. 19M). A third can be recognized, which pos- 
sesses a mosaic of characters, including slightly 
broadened branchiostegal rays along the epihyal, 
and urohyal and pectoral girdle conditions that 
do not conform to either of the above groups. 
Programs REGROUP and WVGM showed little af- 
finity between C. nebulosus of this last group 
and the remainder of the genus. It appears that 
the simple urohyal, broadened rays, and paired 
girdle elements are primitive conditions within 
the Callechelyini, characters shared by C. nebu- 
losus and C. springeri. 


Letharchus Goode and Bean 


Letharchus Goode and Bean 1882: 437. (Type 
species L. velifer Goode and Bean 1882, by 
original designation.) 


DESCRIPTION: Anterior nostril a hole, its rim 
not raised; snout moderate, acute, not rounded 
at tip; median groove on underside of snout 
absent (fig. 32b); intermaxillary and vomerine 
teeth present; DFO above epiotics; anal fin ab- 
sent; four supraorbital pores; neurocranium de- 
pressed, not rounded across parietal-frontal re- 
gion, highest at frontal-parietal suture; hyoid 
arch stout, flexible along CH-EH suture, HH sep- 
arated from CH by a gap; branchiostegal rays 
numerous, slender, all along arch; UH a slender 
filament posteriorly; Hs cartilaginous; pectoral 
girdle contains Cl, SCI, and 2 rod-shaped ele- 
ments; body coloration uniformly dark, con- 
trasting strongly with the white dorsal fin. 


ETYMOLOGY: From the Greek NMNOEOOMAKC 
to forget, and ie or (archos; mascu- 
line), anus, in reference to the lack of an anal fin. 


DISTRIBUTION: Known from three New World 
species: L. velifer from the western Atlantic 


64 


(North Carolina to the northern Gulf of Mexico), 
L. aliculatus McCosker from off Brazil, and L. 
rosenblatti McCosker from the eastern Pacific. 


REMARKS: The genera Letharchus and Paraleth- 
archus were recently revised by McCosker (1974). 
The species of Letharchus form a unique and dis- 
tinctive offshoot from the generalized callech- 
elyin condition in their combination of anal fin 
absence, non-tubular nostrils, an additional supra- 
orbital pore, and the acute snout, differing mark- 
edly from the characters of other genera within 
the tribe. Letharchus pacificus Osborne and 
Nichols and L. opercularis Myers and Wade are 
obviously similar to the above species in lacking 
an anal fin, and probably for that reason were 
assumed by their describers to be congeneric 
with L. velifer. After examining considerable 
material and the types of these five species | 
have concluded that L. opercularis and L. pacifi- 
cus represent a separate generic line within the 
Callechelyini. 

The osteological description of Letharchus is 
based on the eastern Pacific species in that mate- 
rial of L. velifer was unavailable for dissection. 


Leuropharus Rosenblatt and McCosker 


Leuropharus Rosenblatt and McCosker 1970: 502. 
(Type species; L. lasiops Rosenblatt and Mc- 
Cosker 1970, by original designation.) 


DESCRIPTION: Anterior nostril tubular; snout 
short, rounded at tip; median groove on under- 
side of snout absent; snout, nape, and much of 
surface of jaws papillate; intermaxillary teeth 
present, vomerine teeth absent; DFO above SO; 
anal fin present; three supraorbital pores; neuro- 
cranium well rounded, highest anterior to front- 
al-parietal suture; HH separated from CH by a 
gap; branchiostegal rays numerous, slender, and 
along arch; Hs cartilaginous; pectoral girdle con- 
tains Cl, SCl, and 2 rod-shaped elements; body 
coloration nearly uniform, median fins white. 


La 
ETYMOLOGY: From the Greek NE VUPOS 
Ae 
(leuros), smooth, and ACOs (pharos; 
neuter), plow, in reference to the toothless vo- 
mer. 


DISTRIBUTION: A_ single species, known only 
from the type specimen from Manzanillo Bay, 
western Mexico. 


Paraletharchus McCosker 


Paraletharchus McCosker 1974: 620. (Type spe- 
cies; Letharchus pacificus Osburn and Nichols, 


SERIES 4, V.41,#1 McCOSKER — EELS 


1916, by original designation.) 


DESCRIPTION: Anterior nostril tubular; snout 
short, rounded at tip; median groove on under- 
side of snout absent; LL ossicles block-like cyl- 
inders, not heavily fractionated (compare figs. 
22H and 221); intermaxillary and vomerine teeth 
present; GO with a deep anterolateral pocket; 
DFO above SO; dorsal fin elevated; anal fin ab- 
sent; three supraorbital pores; neurocranium well 
rounded, highest anterior to frontal-parietal su- 
ture; hyoid arch stout, only slightly flexible along 
EH-CH suture; HH separated from CH by a nar- 
row gap; branchiostegal rays numerous, along 
arch, distal rays along epihyal broadened basally; 
UH a simple slender filament posteriorly; Hs 
cartilaginous; pectoral girdle contains Cl, SCI, 
and 2 rod-shaped elements; coloration nearly 
uniform to mottled. 


4 
ETYMOLOGY: From the Greek ITAL ERK? 
(para), near, and Letharchus (masculine), a re- 
lated genus. 


DISTRIBUTION: Known from two eastern Pacific 
species, P. opercularis (Myers and Wade), a 
Galapagos endemic, and P. pacificus (Osburn 
and Nichols), ranging from Baja California to 
Costa Rica. 


REMARKS: As mentioned in the remarks on 
Letharchus, the above-mentioned species are not 
congeneric with L. velifer. Schultz and Barton 
(1960) placed L. opercularis in the synonymy of 
L. pacificus, however McCosker (1974) provision- 
ally recognized the Galapagos population as dis- 
tinct on the basis of the difference in the mean 
vertebral number (95% confidence limits: P. 
opercularis 172.7 - 176.6, P. pacificus 158.5 - 
162.3). 


Tribe Sphagebranchini 


TYPE GENUS: Caecula Vahl 1794, the senior ob- 
jective svnonym of Sphagebranchus Bloch, 1795 
(see following remarks). 


DIAGNOSIS: Body (head and trunk) and tail 
moderately elongate, cylindrical, often compres- 
sed posteriorly; body either nearly equal to or 
shorter than tail; snout pointed, often broad and 
depressed; lower jaw included; posterior nostril 
opens into mouth in most genera; GO entirely 
ventral; median fins either very low or absent; 
pectoral fin absent; tail tip sharply pointed; 
head pores well developed, tp? and pop* gener- 
ally, and pop* sometimes present; LL ossicles 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


continuous; teeth conical, not caniniform, and 
generally uniserial; intermaxillary teeth sepa- 
rated by a gap from those of vomer; neurocran- 
ium elongate, generally depressed and truncate 
posteriorly; orbit reduced; otic bulla moderately 
to well developed; hyoid arch generally slender; 
branchiostegal rays few to numerous, generally 
closely associated with hyoid; gill arches re- 
duced, C; absent in most genera; intramuscular 
bones, ribs, and caudal transverse processes well 
developed; precaudal either nearly equal to or 
more numerous than caudal vertebrae; colora- 
tion generally uniform, or darker dorsally. 


REMARKS: Included among the genera of the 
Sphagebranchini are the most frustrating and 
problematical of ophichthid taxa. Their nearly 
complete lack of superficial characters has re- 
sulted in a history of repeated lumping and split- 
ting. Further confusion stems from the inade- 
quate and misleading descriptions of the two 
oldest generic names, Caecula Vahl (1794) and 
Sphagebranchus Bloch (1795). Gosline (1951: 
311) summarized the problem in stating that ‘‘the 
definition, and consequently the limits, of the 
genus Caecula are agreed upon by no two au- 
thors as far as | know.” 

The tribal name Sphagebranchini is derived 
from Swainson’s (1838) family name Sphage- 
branchidae. Swainson’s family name was subse- 
quently rejected (see page 10) long before 
Sphagebranchus Bloch (1795) was shown to be 
a junior objective synonym of Caecula Vahl 
(1794) (see Bohlke and McCosker, 1975). The 
family-group name Caeculidae has not appeared 
in the ichthyological literature. In accordance 
with Article 40 of the International Code of Zoo- 
logical Nomenclature (1964), the family-group 
name Sphagebranchini therefore has priority as 
the tribal name, with Caecula as the type genus 
of the tribe. 


Achirophichthys Bleeker 


Achirophichthys Bleeker 1865: 41. (Type species; 
A. typus Bleeker 1865, by original designation.) 


DESCRIPTION (based on Bleeker, 1865, and 
Weber and de Beaufort, 1916): Body stout, 
slightly longer than tail; snout pointed; eye 
small; anterior nostril not tubular, posterior nos- 
tril opens into mouth; lips with one row of min- 
ute tubercular papillae; DFO slightly behind GO; 
GO low lateral to inferior; teeth conical, long 
and nearly caniniform anteriorly and along vo- 
mer, uniserial on vomer, biserial on maxilla. 


65 


W 
ETYMOLOGY: From the Greek E¢ as 
(achir), without hands, and Ophichthys, (mascu- 
line), the amended spelling of Ophichthus. 


DISTRIBUTION: A single western Pacific species. 


REMARKS: An osteological diagnosis of Achiro- 
phichthys is not included in this study in that 
material of A. typus, the generic type, was un- 
available. A. kampeni (Weber and de Beaufort), 
its sole described congener, is herein referred to 
Lamnostoma. Jordan and Davis (1891: 636) sug- 
gested that A. typus might be the young of 
Brachysomophis crocodilinus, but subsequent 
authors have neither accepted nor commented 
upon this action. Weber and de Beaufort (1916) 
considered Achirophichthys to be a subgenus of 
Brachysomophis. This too was ignored by most 
subsequent authors. Their description of A. 
typus, based on the type specimen, strongly in- 
dicates that it is congeneric with A. kampeni, 
which if true, would place Achirophichthys in 
the synonymy of Lamnostoma. 


Apterichtus Dumeéril 


Caecilia Lacépede 1800: 134. Preoccupied by 
Caecilia Linnaeus, a genus of Amphibia. (Type 
species; C. branderiana Lacépede 1800, by 
monotypy.) 

Apterichtus Duméril 1806: 331. Also spelled 
Apterichthys, Apterichthus, Apterichthe, and 
Apterichtes by other authors. (Type species; 
Muraena caeca Linnaeus 1758, by monotypy.) 

Typhlotes Fischer 1813: 81. A replacement name 
for Caecilia Lacépede, preoccupied. 

Branderius Rafinesque 1815: 93. A replacement 
name for Caecilia Lacépede, preoccupied. 

Ophisurapus Kaup 1856a: 52. (Type species; O. 
gracilis Kaup 1856, by monotypy.) 

Ophisuraphis Kaup 1856b: 29. Emend. pro Oph- 
isurapus Kaup 1856a. 

Verma Jordan and Evermann 1896: 374. (Type 
species; Sphagebranchus kendalli Gilbert 1889, 
by original designation.) 

?Microrhynchus Blache and Bauchot 1972: 728. 
Preoccupied by Microrhynchus Dejean 1821, 
a genus of lepidoptera, as well as mammalia 
(jourdan 1834), crustacea (Bell 1835), aves (Les- 
son 1843) and vermes (Kepner 1935). (Type 
species; Sphagebranchus foresti Cadenat and 
Roux 1964, by original designation.) 


DESCRIPTION: Body very elongate, cylindrical, 
and pointed at both ends; body and tail nearly 
subequal; snout pointed, sub-conical, grooved 


66 


and flattened on underside; lips without barbels; 
eye moderately developed; anterior nostril tubu- 
lar, posterior nostril a horizontally ovate slit out- 
side of mouth; GO ventral, converging forward; 
isthmus short; all fins absent; tp” and pop‘ pres- 
ence variable among species; teeth pointed, uni- 
serial in jaws, and largest at intermaxillary which 
is separated from those of vomer by a short gap; 
skull slightly depressed, sub-truncate to rounded 
posteriorly; orbit reduced; nasals and nasal car- 
tilage developed; SOC short and blunt posteri- 
orly; maxilla pointed posteriorly; opercular mar- 
gins entire; suspensorium anteriorly inclined, 
jaw angle ca. 100°; PG slender, pointed and very 
reduced; hyoid arch slender, HH separated from 
CH by a gap; branchiostegal rays closely asso- 
ciated with hyoid; UH cartilaginous posteriorly; 
C; absent, UP;-UPs separate; Cl broad, SCI re- 
duced, Co and Sc absent; posterior trunk parapo- 
physes with an anterior marginal projection (fig. 
33). 


ETYMOLOGY: From the Greek AMT fov 


(apteron), without fins, and (ichtus, more cor- 
rectly written ichthys; masculine), fish. 


DISTRIBUTION: From 10-12 described and valid 
species, represented in all tropical oceans. 
Bohlke (1968) provisionally reviewed the species 
of Verma (=Apterichtus). The genus Apterichtus 
can be expanded to include: Muraena caeca Lin- 
naeus (M), Caecula gymnocelus Bohlke (EP), C. 
monodi Roux (EA)*, C. equatorialis Myers and 
Wade (EP), Sphagebranchus klanzingai Weber (IP), 
S. flavicaudus Snyder (IP), S. kendalli Gilbert 
(WA), Verma ansp Bohlke (WA), Ophisurapus 
gracilis Kaup  (EA)*, Ophichthys anguiformis 
Peters (EA)*, and possibly Sphagebranchus for- 
esti Cadenat and Roux (EA)* and Microrhynchus 
epinepheli Blache and Bauchot (EA)*. 


REMARKS: Blache and Bauchot (1972) recog- 
nized Verma as distinct from Apterichtus on the 
basis of a minor difference in posterior nostril 
location. Through the kindness of Enrico Tor- 
tonese | have examined a specimen of the Medi- 
terranean Apterichtus caecus, and have con- 
cluded that it is clearly congeneric with the At- 
lantic and Pacific species previously referred to 
Verma. Microrhynchus Blache and_ Bauchot 
(1972) is based upon two species known only 
from the holotypes. Both were unavailable for 
study. Their sketchy description of the external 
morphology of the species, upon which the pres- 
ent study is based, does not provide characters 
which would allow their generic separation from 


SERIES 4, V. 41, #1 McCOSKER — EELS 


Apterichtus, or possibly Ichthyapus. Should Mic- 
rorhynchus prove to be a valid genus a substitute 
name will be required. 


Caecula Vahl 


Caecula Vahl 1794: 149. (Type species; C. ptery- 
gera Vahl 1794, by original designation.) 

Sphagebranchus Bloch 1795: 88. (Type species; 
S. rostratus Bloch 1795 = Caecula pterygera 
Vahl 1794, by monotypy.) 


DESCRIPTION: Body moderately elongate, cylin- 
drical, compressed posteriorly; body and _ tail 
nearly subequal; snout pointed, depressed, and 
broad dorsally, grooved and flattened on under- 
side; eye moderate; anterior nostril flush with 
snout anteriorly, produced as a tube posteriorly, 
posterior nostril associated with a barbel; GO 
entirely ventral, converging anteriorly, much 
longer than isthmus, and with an anterolateral 
duplication forming a deep pouch; DFO slightly 
behind GO; pop* and tp? present, pop* absent; 
teeth conical and uniserial, largest at intermaxil- 
lary which is widely separated from those of 
vomer; skull depressed, broad, and truncate pos- 
teriorly; orbit extremely reduced; nasals and 
nasal cartilage well developed; SOC a short 
broad point posteriorly; maxilla elongate, pointed 
posteriorly; coranoid process of articular moder- 
ately enlarged; operculum well developed, pre- 
operculum narrow and thin, their margins entire; 
suspensorium nearly vertical; hyomandibular 
broad, expanded posterodorsally and anteriorly, 
strongly ridged; otic bulla weakly developed; PG 
slender, pointed at each end, and braced against 
hyomandibular by a posteromedial extension; 
hyoid arch slender; HH separated from CH by 
a gap; branchiostegal rays numerous, flat and 
unbranched, loosely associated with hyoid; out- 
ermost rays along epihyal slightly broadened; 
UH a slender filament posteriorly; C; absent, 
UP3-UPs separated, anterior half of Bi ossified; 
Cl, SCI, Sc and Co present. 


ETYMOLOGY: A diminutive of the Latin caecus, 
blind, regarded as feminine. 


DISTRIBUTION: Two western Pacific species. 


REMARKS: Various species have been haphazard- 
ly assigned to Caecula to such an extent that it 
has become a catch-all for most finned and fin- 
less ophichthids lacking pectoral fins. Smith 
(1964) redescribed the type of C. pterygera and 
began the dissection of this confusing assemb- 
lage. Smith erred in presuming C. pterygera and 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Lamnostoma pictum to be synonymous, and in 
fact, congeneric. Osteologically these genera are 
similar in their coronoid processes, hyoid, gill 
arch, and nostril conditions, but differ tren- 
chently in their neurocrania. 

Blache and Bauchot (1972) placed the type 
species of Sphagebranchus, S. rostratus Bloch, in 
Caecula. BOhlke and McCosker (1975) considered 
the type species of those genera to be conspe- 
cific, and suggested that the collection locality of 
S. rostratus (“river in Surinam’’) was erroneous. 


Cirricaecula Schultz 


Cirricaecula Schultz 1953: 49. Type species; C. 
johnsoni Schultz 1953, by original designa- 
tion.) 


DESCRIPTION: General characters those of /Ich- 
thyapus. Differences include: body and tail near- 
ly subequal; numerous cirri along edge of upper 
lip, a prominent barbel between the nostrils; an- 
terior nostril nearly flush with snout anteriorly, 
slightly produced as a tube posteriorly; GO en- 
tirely ventral, consisting of 2 parallel longitudinal 
slits with thin medial membranes; isthmus min- 
ute; pop’ and tp? present; otic bulla weakly de- 
veloped; PG slender, pointed at each end; hyoid 
arch slender, HH separated from CH by a gap, 
UH a slender filament posteriorly; Cs; ossified, 
UP;-UP, fused; pectoral girdle reduced to a ven- 
trally located Cl pair which are broad and anteri- 
orly expanded (fig. 19); CTP strongly developed. 


ETYMOLOGY: From the Latin cirrus, tendril, and 
Caecula (feminine), a related genus. 


DISTRIBUTION: A single central Pacific species, 
known only from the type series collected at 
Eniwetok, Marshall Islands. 


Hemerorhinus Weber and de Beaufort, 
incertae sedis 


Hemerorhinus Weber and de Beaufort 1916: 280. 
(Type species; Sphagebranchus heyningi Web- 
er 1913, by original designation.) 


DESCRIPTION: Body moderately elongate, cylin- 
drical, pointed at each end; body longer than 
tail; snout pointed, grooved on underside; eye 
small; anterior nostril flush with snout; posterior 
nostril below eye, a long slit in upper lip; GO 
inferior, vertical; vertical fins low; tp? present(?), 
pop® absent(?). 


/ 
ETYMOLOGY: Presumably from the Greek 72 ME 
efi (hemeros), cultivated, and eC C EI 6 


67 


(rhinos; masculine in accordance with item 
30(a)(3) of the International Code of Zoological 
Nomenclature), nose. 


DISTRIBUTION: Known from the type specimen, 
collected in deep water (69-91 meters) from 
Flores, Indonesia, and H. opici Blache and Bau- 
chot (EA)*. 


REMARKS: Species of Hemerorhinus were un- 
available for study. Blache and Bauchot’s (1972) 
redescription of the type species clarified several 
confusing aspects of the original description, par- 
ticularly in correcting the mistaken intepretation 
of the nostrils and fin position. From their dis- 
cussion however, | am unable to confidently find 
its placement within this tribe. On the basis of 
Blache and Bauchot'’s illustrations, the cephalic 
pore condition would indicate a similarity to the 
species of Yirrkala. 


Ichthyapus de Barneville 


Ichthyapus de Barneville 1847: 219. (Type spe- 
cies; /. acutirostris de Barneville 1847, by 
monotypy.) 

Rhinenchelys Blache and Bauchot 1972: 718. 
(Type species; Sphagebranchus ophioneus 
Evermann and Marsh 1902, by original desig- 
nation.) 


DESCRIPTION: Body elongate, cylindrical, point- 
ed at both ends; tail longer than body; snout 
pointed, depressed, and broad dorsally, grooved 
and flattened on underside; lips without barbels; 
eye small; anterior nostril flush with snout, pos- 
terior opens into mouth; GO entirely ventral, 
with a thin medial membrane, converging for- 
ward, isthmus small; all fins absent; tp? always 
and pop? usually present; teeth pointed, uni- 
serial, and largest at intermaxillary which are 
separated from those of vomer by a gap; skull 
depressed, broad, and truncate posteriorly (fig. 
8); orbit extremely reduced; nasals moderately, 
and nasal cartilage well developed; SOC moder- 
ately projecting posteriorly; maxilla elongate and 
pointed posteriorly; suspensorium nearly verti- 
cal; opercular margins entire, preopercle re- 
duced; hyomandibular broad, expanded anteri- 
orly and posterodorsally; otic bulla moderately 
developed; PG elongate and rectangular posteri- 
orly, with a slender projection from the antero- 
dorsal corner; hyoid thickened (not as slender as 
in related genera); HH separated from CH by a 
narrow gap; branchiostegal rays not numerous, 
slender and generally unbranched, closely asso- 


68 


ciated with hyoid, outermost rays along EH 
broadened basally; UH cartilaginous for pos- 
terior two-thirds; C; reduced (see Remarks), UP3- 
UP, separate; Cl broad, SCI usualy absent, Sc 
and Co absent; posterior trunk parapophyses 
with an anterior marginal projection as in Apter- 
ichtus (fig. 33). 


ETYMOLOGY: From the Greek ¢Y% Obs 
(ichthys), fish, and Gx ae) Sp (apous; 
masculine), without foot, presumably in _refer- 
ence to the lack of pectoral fins. 


DISTRIBUTION: Circumtropical, with 6-7 valid 
species. Included are: /. acutirostris de Barneville 
(locality unknown)*, Sphagebranchus — vulturis 
Weber and de Beaufort (=Caecula platyrhyncha 
Gosline) (IP), S. ophioneus Evermann and Marsh 
(WA), Apterichthys selachops Jordan and Gilbert 
(EP), and three undescribed eastern and western 
Pacific forms. Incertae sedis: Sphagebranchus 
omanensis Norman (IP). 


REMARKS: The sharp-snouted finless species pre- 
viously placed in Sphagebranchus, excluding the 
species of Cirricaecula and Apterichtus as herein 
defined, are referred to Ichthyapus. The generic 
type, /. acutirostris, is obviously congeneric with 
those species according to descriptions of the 
type made by de Barneville (1847, fide Fowler, 
1936: 293), Kaup (1856b: 29) and Blache and 
Bauchot (1972: 718-728). 

Blache and Bauchot (1972) differentiated 
Rhinenchelys from Ichthyapus on the basis of 
minor differences in nostril condition and inter- 
maxillary tooth location. My examination of the 
osteology of ophioneus, the type of Rhinench- 
elys, indicates that it is congeneric with sela- 
chops, vulturis, and presumably acutirostris. 

Nelson (1966a: table 1, figure 19) has de- 
scribed and illustrated the gill arch condition of 
I. vulturis (as Caecuia platyrhyncha). The fifth 
ceratobranchial (Cs) is reduced and fused to the 
lower pharyngeal dermal tooth plate. | have 
found the gill arches of J. ophioneus, |. vulturis, 
and /. selachops to be similar in this condition. 
Cirricaecula, with a prominent Cs, appears inter- 
mediate between /Ichthyapus and most Ophich- 
thyini in this condition. The C; condition of the 
related genus Apterichtus, based on my examina- 
tion of A. flavicaudus, is the most reduced in 
the group. 


Lamnostoma Kaup 


Lamnostoma Kaup 1856: 49 (23). (Type species; 
L. pictum Kaup 1856 = Dalophis orientalis 


SERIES 4, V.41,#1 McCOSKER — EELS 


McClelland 1844, by Jordan 1919b as first re- 
viser.) 

Anguisurus Kaup 1856: 50 (24). (Type species; 
A. punctulatus Kaup 1856 = Dalophis orien- 
talis McClelland 1844, by monotypy.) 


DESCRIPTION: Body stout, cylindrical, pointed 
at each end; body slightly longer than tail; snout 
pointed, its underside grooved; eye small to 
moderate; anterior nostril flush along snout, its 
posterior rim produced, posterior nostril usually 
associated with a pendulous flap; GO inferior, 
ca. equal to isthmus; DFO above or behind GO; 
tp? and pop? absent; teeth slender, pointed, and 
recurved, uniserial or biserial in jaws, those of 
intermaxillary and vomer largest and _ widely 
spaced; neurocranium truncate posteriorly, 
elongate and narrow, particularly along ethmoid 
and interorbital region; orbit depressed; nasals 
and nasal cartilage moderately developed; SOC 
present; maxillae moderately elongate, slender 
but not pointed posteriorly; coronoid process of 
articular greatly enlarged; opercular series mod- 
erately developed, their margins entire; suspen- 
sorium anteriorly inclined, jaw angle ca. 100°; 
hyomandibular broad, expanded posterodorsally 
and strongly ridged; otic bulla well developed; 
PG slender, very reduced; hyoid slender, equal 
to branchiostegal rays in thickness, HH separated 
from CH by a broad gap, rays numerous, flat and 
unbranched, only the distal-most associated with 
hyoid, others terminate anteriorly behind tip of 
slender UH, outermost rays along EH slightly 
broadened; gill arches reduced, C; absent, UPs- 
UP, separate, B; cartilaginous except at anterior 
tip which is ossified; Cl, SCI, and reduced Co 
and Sc present; coloration generally darker dor- 
sally, a series of white spots across nape. 


ETYMOLOGY: From the Greek AMX AL CX 

(lamna), a horrible anthropophagous monster, a 
bugbear used by the Greeks to frighten refract- 
ory children (Jordan and Evermann, 1896: 49), 
and g TO (stoma; neuter), mouth. 


DISTRIBUTION: From the western Pacific, pro- 
visionally including four species. 


REMARKS: Lamnostoma has been placed by re- 
cent authors in the synonymy of Caecula. It is 
herein found to differ markedly in numerous 
osteological and morphological characters, and 
is COnsequently resurrected. The species of this 
genus are generally collected in freshwater, and 
are easily recognized by their slender jaws and 
conspicuous white spotting on the head and 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


lateral line. Several nominal species are included 
in Lamnostoma, most of which fall into the syn- 
onymy of L. orientalis (McClelland). Caecula min- 
dora Jordan and Richardson and C. tay/ori Herre 
are also referable to this genus. Achirophichthys 
kampeni (Weber and de Beaufort) is clearly a 
Lamnostoma. Its description, and recent refer- 
ences containing general morphological descrip- 
tions (Herre, 1924; Nichols, 1955; La Monte, 
1961; Tortonese, 1964), concern large adults 
which have been collected in freshwater, each 
displaying the slender jaws and the head and 
body spotting of Lamnostoma. 


Stictorhinus Bohlke and McCosker 


Stictorhinus Bohlke and McCosker 1975: 5. 
(Type species; S$. potamius Bohlke and McCos- 
ker 1975, by original designation.) 


DESCRIPTION: General characteristics those of 
Ichthyapus. Differences include: Median fins 
present, low but distinct, DFO ca. mid-trunk; 
tp? present, pop* absent; LL ossicles with a short 
gap at pores; basisphenoid not elongate as in 
Ichthyapus (compare Figs. 8 and 9); SOC re- 
duced; opercular series reduced, interopercle 
absent, preopercle reduced; branchiostegal rays 
numerous, branched basally in some individuals, 
closely associated with hyoid, outermost rays 
along EH broadened basally; UH a slender ossi- 
fied filament posteriorly; C; absent; Cl broad 
and expanded anteriorly, SCI absent, Sc and Co 
reduced; trunk parapophyses lack an anterior 
projection (fig. 33). 


ETYMOLOGY: From the Greek TTLKTOS 
(stiktos), pricked or punctured, and eC Llp 

(rhin, latinized to rhinus, nominative case), nose, 
in reference to the nature of the anterior nostrils. 


DISTRIBUTION: A single western Atlantic species, 
known from tidal rivers in Brazil. 


REMARKS: In 1971 Naercio Menezes sent the 
author a series of eels from freshwater in Brazil. 
These were identified as Sphagebranchus rostra- 
tus Bloch, a species known only from the type 
specimen said to have been from a “river in Suri- 
nam.”” Subsequent examination of the type speci- 
mens of Caecula pterygera and Sphagebranchus 
rostratus by J. E. BOhlke found them to be based 
on the same species, and the Brazilian specimens 
to be an undescribed genus and species (Bohlke 
and McCosker, 1975). My treatment (McCosker, 
1973) of Sphagebranchus was therefore based on 
Stictorhinus. 


69 


Yirrkala Whitley 


Yirrkala Whitley 1940: 410. (Type species; Y. 
chaselingi Whitley 1940 = Sphagebranchus 
lumbricoides Bleeker 1865, by original desig- 
nation.) 

Pantonora Smith 1964: 719. (Type species; Oph- 
ichthys tenuis Gunther 1870, by original desig- 
nation.) 


DESCRIPTION: Body elongate, cylindrical, shorter 
than tail; snout conical, moderately developed, 
its underside grooved; eye moderate; anterior 
nostril in a short tube, posterior opens into 
mouth; GO ventral, longer than isthmus; DFO 
above or slightly behind GO; tp” usually present, 
pop® absent; teeth conical, pointed, nearly sub- 
equal, uniserial in jaws, those on vomer biserial 
anteriorly, separated from those of intermaxillary 
by a gap; skull not depressed, rounded posteri- 
orly; orbit not strongly depressed; nasals and 
nasal cartilage moderately developed; SOC ab- 
sent; maxilla pointed posteriorly; opercular ser- 
ies well developed, their margins entire; sus- 
pensorium anteriorly inclined, jaw angle ca. 
100°; PG slender, pointed anteriorly; HH sepa- 
rated from CH by a short gap (HH absent in Y. 
misolensis); branchiostegal rays closely associ- 
ated with hyoid; UH with a short projection pos- 
teriorly; C; absent, UP3-UPs separate; pectoral 
girdle contains SCI, Cl, and reduced Co and Sc; 
posterior trunk parapophyses lack anterior mar- 
ginal projections. 


ETYMOLOGY: Named for Yirrkala, northern Aus- 
tralia, the type locality of the type species, mas- 
culine, in accordance with item 30(b)(ii) of the 
International Code of Zoological Nomenclature. 


DISTRIBUTION: Contains approximately 12 spe- 
cies, from the eastern Atlantic, the Red Sea, and 
Indian and western Pacific Oceans. Included are: 
Sphagebranchus lumbricoides Bleeker, Ophich- 
thys tenuis Gunther, Caecula maculata Klause- 
witz, and an undescribed species from the Mar- 
quesas. Incertae sedis: Sphagebranchus macro- 
don Bleeker*, S. gjellerupi Weber and de Beau- 
fort*, S. kaupi Bleeker, Dalophis moluccensis 
Bleeker*, Ophichthys misolensis Gunther, Mur- 
aena fusca Zuiew*, and Caecula natalensis Fow- 
ler*. 


REMARKS: The description of Yirrkala is so lack- 
ing in diagnostic characters as to preclude its 
proper placement, and consequently has been 
ignored by most recent authors. The only sub- 
sequent reference to Y. chaselingi, the generic 


70 


type, which | have encountered is that of Munro 
(1957) in which this species is placed in Sphage- 
branchus. | have examined and x-rayed the badly 
damaged and dessicated paratype of Y. chase- 
lingi. The anterior nostrils, head pores, and gill 
openings are too badly damaged to allow accur- 
ate examination, however the dentition is well 
preserved and agrees well with Whitley’s illus- 
tration. In his description, Whitley stated that 
Y. chaselingi and Sphagebranchus lumbricoides 
are congeneric, differing in the ‘position of 
dorsal origin, proportion of head to trunk, in 
having vomerine teeth largest, and other details 
of porportions and dentition.” My comparison of 
the paratype of Y. chaselingi and specimens of 
S. lumbricoides from Vietnam (CAS 13969) and 
the Philippines (CAS reg. 1607) indicates that 
they are conspecific. The proportions, dorsal fin 
origins, dentition, and vertebral numbers (Y. 
chaselingi = 153, S. lumbricoides = 151,154) 
are not different. 

Pantonora Smith (1964) is herein considered a 
synonym of Yirrkala. 

Sphagebranchus kaupi and Ophichthys misol- 
ensis are provisionally referred to Yirrkala. Y. 
kaupi differs in having the major axis of its GO 
vertical. Y. misolensis differs in having basihyals 
fused to the ceratohyals, lateral line ossicles con- 
sisting of numerous short coils rather than small 
block-like segments, and lacking tp. 

The external morphology of the species of 
Hemerorhinus, as described by Blache and Bau- 
chot (1972), indicates that those species might 
be conspecific with the species within this com- 
plex. A comparative study of the type species of 
Yirrkala, Hemerorhinus, and Pantonora is clearly 
needed. 


Tribe Bascanichthyini 


TYPE GENUS: Bascanichthys Jordan and Davis, 
1892. 


DIAGNOSIS: Body (head and trunk) and tail 
moderately to extremely elongate, generally cyl- 
indrical, and compressed posteriorly in some 
genera; body either equal to or longer than tail; 
lower jaw included; posterior nostril opens into 
mouth; GO low lateral, crescentric, never en- 
tirely ventral; median fins generally low, DFO 
on head in most genera; pectoral fin absent or 
present as a minute flap in upper GO corner; 
head pores reduced, pop* and tp? absent; LL 
ossicles nearly continuous or separated by a 
short gap at pores; teeth conical, not caniniform; 


SERIES 4, V.41,#1 McCOSKER — EELS 


neurocranium variable in proportions and pos- 
terior shape; orbit generally reduced; otic bulla 
moderately to well developed; hyoid condition 
variable; branchiostegal rays numerous, associ- 
ated with the hyoid; gill arches reduced, Cs; ab- 
sent in most genera; IM bones, ribs, and CTP 
developed; precaudal vertebrae generally more 
numerous than caudal; coloration nearly uni- 
form, or darker posteriorly. 


Allips McCosker 


Allips McCosker 1972: 116. (Type species; A. 
concolor McCosker 1972, by original designa- 
tion.) 


DESCRIPTION: Body elongate, cylindrical for 
most of its length; body much longer than tail; 
head markedly rugose; snout blunt, its under- 
side grooved; eye small; anterior nostril tubular; 
GO oblique, lateral, and less than isthmus in 
length; median fins low, DFO in anterior trunk 
region; pectoral minute; caudal blunt; tip of 
snout with numerous sensory hairs; LL ossicles 
nearly continuous; teeth small, conical, uniserial, 
intermaxillary teeth largest, separated from those 
of vomer; skull sloping posteriorly, orbit a 
narrow slit; SOC weakly developed; maxillae 
taper posteriorly; HH separated from CH by a 
gap; branchiostegal rays numerous, slender; pec- 
toral girdle reduced, only Cl, SCI, and a thin 
Co (2). 


yA 
ETYMOLOGY: From the Greek & VAOS 
(allos), another, and oe yy (ips; mascu- 
line), a worm. 


DISTRIBUTION: A_ single species, known only 
from the type specimen from Thailand. 


Bascanichthys Jordan and Davis 


Bascanichthys Jordan and Davis 1891: 621. (Type 
species; Caecula bascanium Jordan 1885 = 
Sphagebranchus teres Goode and Bean 1882, 
by original designation.) 


DESCRIPTION: Body elongate, cylindrical, and 
compressed posteriorly; body longer than tail; 
head markedly rugose; snout short and blunt, 
its underside grooved; mouth small; eye small; 
anterior nostril tubular; GO nearly horizontal, 
low lateral, = isthmus in length; median fins 
low, DFO on head; pectoral minute; caudal 
blunt; LL ossicles nearly continuous; teeth small, 
conical, uniserial in jaws, intermaxillary teeth 
separated from those of vomer by a short gap; 
neurocranium sloping posteriorly (fig. 10), orbit 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


reduced; nasals and nasal cartilage moderately 
developed, the nasals closely associated with the 
ethmoid for their entire length; SOC absent; 
maxilla weak, slender, and pointed posteriorly, 
attached before mid-vomer (fig. 16); opercular 
series moderately developed, their margins en- 
tire; suspensorium nearly vertical; otic bulla 
moderately developed; PG pointed anteriorly; 
HH separated from CH by a narrow suture; 
branchiostegal rays numerous, often branched 
basally, all along arch, the distal rays not broad- 
ened basally; UH either ossified or cartilaginous 
posteriorly; C; reduced or absent, UPs3-UPs sep- 
arate; pectoral girdle consists of reduced Cl, 
SCI, Sc and Co. 


ETYMOLOGY: From Bascanion (diminutive, from 
the Greek 32a 7K X vos , malignant), 
the generic name for the black snake, and 

oe e&f (ichthys; masculine), fish. 


DISTRIBUTION: All tropical oceans, with ap- 
proximately 16 presently recognized species, in- 
cluding an undescribed eastern Pacific species 
from Cocos Island. Included are: B. bascanoides 
Osburn and Nichols (EP), B. cylindricus Meek and 
Hildebrand (EP), B. panamensis Meek and Hilde- 
brand (EP), B. ceciliae Blache and Cadenat (EA)*, 
B. congoensis Blache and Cadenat (EA)*, B. paul- 
ensis Storey (WA), B. pusillus Seale (IP)*, Sphage- 
branchus teres Goode and Bean (WA), S. longi- 
pinnis Kner and Steindachner (IP)*, S. scuticaris 
Goode and Bean (WA)*, Callechelys myersi Herre 
(IP), Ophichthys filaria Gunther (IP)*. I/Incertae 
sedis: Callechelys longissimus Cadenat and Mar- 
chal (EA)*, and Ophichthys kirkii Gunther (IP)*. 
Leptenchelys tenuis Tortonese, from New Guinea, 
is probably a junior synonym of B. longipinnis. 


REMARKS: The species of Bascanichthys have 
been reviewed by Storey (1939), Ginsburg (1951) 
(western Atlantic), and Blache and Cadenat (1971) 
(eastern Atlantic). The status of Bascanichthys 
pusillus, considered a junior synonym of B. filaria 
by Fowler (1931: 316), is uncertain. 


Caralophia Bohlke 


Caralophia Bohlke 1955: 1. (Type species; C. 
loxochila Bohlke 1955, by original designa- 
tion.) 


DESCRIPTION: Body elongate, cylindrical, mod- 
erately compressed posteriorly; body longer than 
tail; snout subconical, blunt from above, lacking 
a groove on its underside; eye moderate; anter- 
ior nostril non-tubular, a hole with lateral projec- 


ail 


tions into it; GO ventral, converging forward, 
longer than isthmus; median fins low, DFO on 
head; tail tip pointed; LL narrowly separted at 
pores; teeth bluntly conical, uniserial in jaws 
and on vomer, intermaxillary teeth slightly larger, 
separated from those of vomer by a short gap; 
skull sloping posteriorly, depressed dorsally, or- 
bit reduced; nasals stout and well developed, 
their margins entire, tightly joined to ethmoid; 
nasal cartilage well developed; frontals extend 
posteriorly to mid-parietals; SOC moderately de- 
veloped, SO extends from mid-frontals posteri- 
orly to a strong point; maxilla weak, slender, and 
pointed posteriorly; opercular series developed, 
their margins entire; suspensorium posteriorly 
inclined, jaw angle ca. 100°; otic bulla moder- 
ately developed; PG broad centrally, closely as- 
sociated with maxilla anteriorly; hyoid stout, HH 
fused to CH; branchiostegal rays numerous, 
along arch, and often branched basally; distal 
rays along EH broadened basally; UH reduced to 
a well ossified basal plate with a posterior car- 
tilaginous filament; tooth plates reduced, UP3-UP4 
fused; pectoral girdle reduced to a Cl and frag- 
ments of a SCI, Sc, and Co. 


ETYMOLOGY: From the Greek KX PK , head, 
and NAO Pf cot (lophia; feminine), a mane, 
in reference to the anterior dorsal fin origin. 


DISTRIBUTION: A single western Atlantic spe- 
cies, extending from the Bahamas to the lesser 
Antilles and Panama. 


Dalophis Rafinesque 


Dalophis Rafinesque 1810a: 68. (Type species; 
D. serpa Rafinesque 1810a = Sphagebranchus 
imberbis De la Roche 1809.) 

Pterurus Rafinesque 1810b: 59. (Type species; 
P. flexosus Rafinesque 1810b = Sphagebranch- 
us imberbis De la Roche 1809, by monotypy.) 

Scytallurus Duméril 1856: 199. (Type species; 
Sphagebranchus imberbis De la Roche 1809, 
by monotypy.) 

Pelia Bleeker 1863: 128. (Type species; P. cepha- 
lopeltis Bleeker 1863.) 


DESCRIPTION: Body moderately elongate, cyl- 
indrical, slightly compressed posteriorly; _ tail 
longer than body; snout sub-conical, grooved 
on underside; eye small; anterior nostril tubular; 
GO low on body, extending onto venter, its 
length ~ isthmus; vertical fins low, DFO well 
behind GO; pectoral fin rudimentary if present; 
tail tip blunt; LL ossicles widely separated at 


72 


pores; teeth conical, uniserial, intermaxillary 
teeth separated from those of vomer by a gap; 
skull subtruncate posteriorly, orbit depressed; 
nasals well developed; SOC weakly developed; 
maxilla tapers posteriorly; suspensorium nearly 
vertical; HH separated from CH by a gap; UH 
ossified posteriorly; gill arches stout, Hs ossified, 
C; a slender ossified rod, UP3-UPs separate; pec- 
toral girdle reduced, only a Cl and SCI present. 


ETYMOLOGY: The significance of the prefix is 
not obvious, but possibly derived from Dalmatia, 
then a country on the east side of the Adriatic 
Sea, and ro) na (ophis; masculine), a 
snake. 


DISTRIBUTION: Five eastern Atlantic and Medi- 
terranean species are presently recognized, in- 
cluding: D. boulengeri Blache and Bauchot (EA)*, 
D. multidentatus Blache and Bauchot (EA)*, D. 
obtusirostris Blache and Bauchot (EA)*, Pelia 
cephalopeltis Bleeker (EA)*, and Sphagebranchus 
imberbis De la Roche (M). 


REMARKS: Blache and Bauchot (1972) have re- 
cently expanded Dalophis to include Pelia and 
five nominal species. Their finding (p. 746) that 
the pectoral fin, though generally absent, may be 
represented by a ‘miniscule filament’ is in 
agreement with Lozano Rey’s (1947, p. 546). 
Through the kindness of Enrico Tortonese | have 
been able to examine and partially dissect a 
specimen of Dalophis imberbis. A complete os- 
teological preparation, however, was impossible. 

Dalophis has been summarily synonymized 
with Caecula by previous authors. Its general 
facies (physiognomy, low median fins, and body 
depth and taper) and certain osteological fea- 
tures (dentition, pectoral girdle, and anterior trunk 
vertebrae) suggest a relationship with Ethadophis 
and Leptenchelys, genera restricted to the east- 
ern Pacific. The gill arch skeleton and body/tail 
proportions are typically ophichthin, and for that 
reason, Dalophis is placed in the Bascanichthyini 
with reservations. 


Ethadophis Rosenblatt and McCosker 


Ethadophis Rosenblatt and McCosker 1970: 498. 
(Type species; EF. byrnei Rosenblatt and Mc- 
Cosker 1970, by original designation.) 


DESCRIPTION: Body moderately elongate, cyl- 
indrical anteriorly, becoming compressed pos- 
teriorly; body equal to or slightly longer than 
tail; snout rounded, conical from above; eye 
small; anterior nostril tubular; GO low on body, 


SERIES 4, V. 41, #1 McCOSKER — EELS 


extending onto venter, their length = to isthmus; 
vertical fins low, DFO before GO; tail tip blunt, 
fleshy in one species; LL ossicles separated at 
pores; teeth conical, uniserial, intermaxillary 
teeth separated from those of vomer by a gap; 
skull subtruncate posteriorly, orbit depressed; 
nasals well developed; SOC moderately devel- 
oped, pointed as in Bascanichthys; maxilla tap- 
ers posteriorly; suspensorium nearly vertical; HH 
separated from CH by a gap; branchiostegal rays 
numerous, slender and along hyoid; UH ossified 
posteriorly; UP3-UP,s separate; pectoral girdle re- 
duced, only a Cl, SCI, and Co (?) remnant visible 
by radiograph. 


Zz 
ETYMOLOGY: From the Greek cen 
(ethas), customary or ordinary, and O PoG 
(ophis; masculine), serpent. 


DISTRIBUTION: Two species, £. byrnei Rosen- 
blatt and McCosker and £. merenda Rosenblatt 
and McCosker, known only from the type speci- 
mens from Baja California, Mexico. 


Gordiichthys Jordan and Davis 


Gordiichthys Jordan and Davis 1891: 644 (Type 
species; G. irretitus Jordan and Davis 1891, 
by original designation.) 


DESCRIPTION: Body extremely elongate, cylin- 
drical, much longer than tail; snout tapering; eye 
moderate; GO low lateral; DFO behind nape: 
teeth conical, recurved, and uniserial. 


ETYMOLOGY: From  Gordius, the horsehair 
worm, named after I-76 P Alos, the king 
whose complicated knot was cut by Alexander, 
and ox evr (ichthys; masculine), fish. 


DISTRIBUTION: Known from a single deep-water 
western Atlantic species. 


REMARKS: The above description is based on 
Jordan and Davis (1891) and Ginsburg (1951) in 
that material of G. irretitus, the type species, was 
unavailable for study. Gordiichthys will be rede- 
scribed by J. E. Bohlke (personal communication). 
G. springeri Ginsburg, its sole described con- 
gener, is a species of Callechelys (fide Rosenblatt 
and McCosker, 1970, and McCosker and Rosen- 
blatt, 1972). 


Leptenchelys Myers and Wade 


Leptenchelys Myers and Wade 1941: 72. (Type 
species; L. vermiformis Myers and Wade 1941, 
by original designation.) 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


DESCRIPTION: Body elongate, cylindrical, slightly 
compressed posteriorly; body longer than tail; 
snout short, subconical; top of head and snout 
covered with numerous papillae; eye small; an- 
terior nostril tubular; GO low lateral, crescentic, 
= isthmus; DFO on head; median fins conflu- 
ent around tail tip; teeth weak, pointed, uni- 
serial; skull sloping posteriorly, orbit reduced; 
SO rounded, not projecting posteriorly; suspen- 
sorium anteriorly inclined, jaw angle ca. 100°; 
hyoid arch slender, HH separated from CH by a 
narrow gap: branchiostegal rays slender, appear 
from radiograph to lie along hyoid; pectoral 
girdle reduced, only SCI and Cl visible in radio- 
graph. 


ETYMOLOGY: From the Greek NEMTTOS 
(leptos), thin, and é YK ENS (enchelys; 
either feminine or masculine, treated as mascu- 
line), eel. 


DISTRIBUTION: A _ single species, known only 
from the type specimen from Playa Blanca, 
Pacific Costa Rica. 


REMARKS: This poorly known genus is provision- 
ally referred to the Bascanichthyini. The generic 
type, apparently uncollected since the capture 
of the 115 mm type specimen, was examined and 
radiographed for the purpose of this study. Its 
inclusion as a bascanichthyin is based on the 
numerous branchiostegal rays which appear to 
contact the hyoid, the posteriorly sloping neuro- 
Cranium, the low unconstricted gill openings, 
and the head pore configuration. Caudal fin rays 
are present on the type specimen, but are more 
poorly developed than any myrophine’s. 

Leptenchelys has had an erratic history. Origi- 
nally placed in the Echelidae on the basis of its 
having a caudal fin (Myers and Wade, 1941), it 
was then synonymized with Muraenichthys by 
Schultz and Woods (1949), re-erected and ex- 
panded by Schultz, et al. (1953), and finally, re- 
stricted to a single species (McCosker, 1970) 
differing markedly from Muraenichthys and re- 
lated genera. Leptenchelys tenuis Tortonese 
(1964) from New Guinea is a species of Bascan- 
ichthys, which is possibly synonymous with B. 
longipinnis described from Samoa. My exami- 
nation of the holotype of L. tenuis indicates that 
Tortonese overlooked the rudimentary pectoral 
fin, similarly developed in B. longipinnis accord- 
ing to Storey (1939). The tail tip of L. tenuis is 
soft and fleshy, but not unlike that of other 
species of Bascanichthys. 


7k) 


Phaenomonas Myers and Wade 


Phaenomonas Myers and Wade 1941: 77. (Type 
species; P. pinnata Myers and Wade 1941, by 
original designation.) 


DESCRIPTION: Body elongate, cylindrical, much 
longer than tail; head markedly rugose; snout 
blunt, grooved ventrally; eye minute; anterior 
nostril tubular; GO low lateral, oblique and 


elongate, = isthmus; DFO mid-head, low, end- 
ing in anterior trunk region; anal fin absent; 
teeth small, conical, uniserial, intermaxillary 


teeth largest, separated from those of vomer; 
skull sloping posteriorly, orbital foramen a_nar- 
row slit; nasal cartilage weak; SOC weakly de- 
veloped; maxilla tapers potseriorly; opercular 
series reduced, all but opercle serrated at mar- 
gin; suspensorium anteriorly inclined, jaw angle 
ca. 100°; otic bulla well developed; PG free and 
tapering anteriorly; HH separated from CH by a 
gap; branchiostegal rays numerous, slender and 
along hyoid; UH club-shaped, cartilaginous pos- 
teriorly; pectoral girdle reduced, only SCI, Cl, 
and a thin Co (?); IM bones, ribs, and CTP 
weakly developed. 


ETYMOLOGY: Presumably from the Greek 
PAC Ya>(phainos), to show, and Lo va 
(monas; feminine), single or alone, in reference 
to the unique dorsal fin condition. 


DISTRIBUTION: Two described species, P. pin- 
nata Myers and Wade ranging from the Gulf of 
California to Colombia, and P. cooperae Palmer, 
ranging from Hawaii to the western Indian 
Ocean (McCosker, In Press). 


Tribe Ophichthini 
TYPE GENUS: Ophichthus Ahl, 1789 


DIAGNOSIS: Body (head and trunk) and _ tail 
moderately elongate, cylindrical anteriorly, gen- 
erally compressed posteriorly; body usually 
shorter than tail; snout conical or subconical; 
lower jaw usually included; anterior nostril gen- 
erally tubular; posterior nostril opens into mouth 
in most genera; gill openings lateral, their major 
axis vertical, crescentic, about equal to, less than, 
or longer than isthmus; median and pectoral fins 
present; tail tip sharply to bluntly pointed; tp? 
and pop‘ absent; dentition variable, including 
caniniform and molariform conditions; neuro- 
cranium moderately elongate; orbit well devel- 
oped; otic bulla moderately to well developed; 
hyomandibular moderately to strongly ridged; 


74 


opercular series well developed, margins entire; 
gill arches well developed, C; present in many 
genera; pectoral girdle generally well developed; 
intramuscular bones, ribs, and caudal transverse 
processes well developed; coloration variable. 


Aplatophis Bohlke 


Aplatophis Bohlke 1956b: 1. (Type species; A. 
chauliodus Bohlke 1956b, by original designa- 
tion.) 


DESCRIPTION: Body stout, not elongate, later- 
ally compressed posteriorly; body slightly longer 
than tail; jaws elongate, the lower projecting be- 
yond the upper; eye small; posterior nostril in 
a short tube before and beneath eye; GO verti- 
cal, lateral and elongate, less than isthmus; DFO 
behind pectoral tips; tail tip blunt; pop* absent, 
median head pores absent although frontal com- 
missure and supratemporal canal are present; LL 
ossicles widely separated at pores; anterior teeth 
of both jaws excessively developed as long fang- 
like canines extending far outside mouth when 
closed, jaw teeth biserial, those of vomer few 
and uniserial and continuous with those of inter- 
maxillary; skull subtruncate posteriorly, orbit 
large, eyeball displaced anteriorly; PO ossicles 
weak, not fused as a strut to maxilla; nasals ex- 
tremely reduced to a short and slender canal, 
nasal cartilage absent; frontal midline elevated 
anteriorly forming a sharp ridge extending to 
‘ $O; SOC absent, SO anteriorly contacts frontals 
and separates parietals; maxilla elongate, round- 
ed posteriorly; opercular series developed, sub- 
opercle scythe-shaped, preopercle enlarged; sus- 
pensorium posteriorly inclined, jaw angle ca. 
80°; hyomandibular ridged and elongate pos- 
terodorsally for muscle attachment; otic bulla 
well developed; PG broad posteriorly, free and 
terete anteriorly, HH separated from CH by a 
gap; branchiostegal rays slender, rays of EH 
joined basally; UH a spike posteriorly; Cs ossi- 
fied, UPs-UP,s separate, Bo-, absent; pectoral fin 
moderately developed, girdle well developed, 
Sc and Co large, actinosts absent (fig. 19F); cau- 
dal vertebrae fewer than precaudal; coloration 
nearly uniform, slightly darker dorsally. 


Ya 
ETYMOLOGY: From the Greek XT AAT OS 
(aplatos), unapproachable, and S PSS (ophis; 
masculine), serpent. 


DISTRIBUTION: Known from a single trans-At- 
lantic species. 


SERIES 4, V.41,#1 McCOSKER — EELS 


Brachysomophis Kaup 


Brachysomophis Kaup 1856: 45 (9). (Type spe- 
cies; B. horridus Kaup 1856 = Ophisurus croc- 
odilinus Bennett 1833, by monotypy.) 


DESCRIPTION: Body cylindrical, moderately 
elongate, longer than tail; snout very short, flat, 
blunt; jaws elongate, with lower jaw slightly pro- 
truding; eye moderate; a conspicuous transverse 
depression exists in the postorbital region of 
some species; lips with numerous small cirri; an- 
terior nostril in a very short tube, posterior opens 
into mouth; GO low lateral, crescentic, longer 
than isthmus; DFO well behind pectoral tips; tail 
tip sharply pointed; pop* absent; LL ossicles 
separated at pores; teeth strong, conical, maxil- 
lary biserial, dentary uniserial, those of vomer 
uniserial, large, and widely separated anteriorly 
but continuous with those of intermaxillary; skull 
subtruncate posteriorly, orbit reduced; dorsal- 
most postorbital with a broad anterior flange 
forming posterior margin of orbit, tightly sutured 
to smaller postorbitals to form a maxillary strut 
(fig. 15); nasals and nasal cartilage much re- 
duced; SOC absent, SO broad posteriorly; max- 
illa elongate, tapering posteriorly; opercular 
series well developed, margins slightly serrate; 
suspensorium nearly vertical; hyomandibular 
ridged; otic bulla well developed; PG broad 
posteriorly, tapering to a fine point anteriorly; 
hyoid arch slender, HH separated from CH by 
a gap; branchiostegals slender and unbranched, 
closely associated with hyoid; UH a spike pos- 
teriorly; Cs ossified, UPs3-UPs separate; pectoral 
fin and girdle reduced, SCI and actinosts absent, 
SC reduced; caudal vertebrae fewer than pre- 
caudal; coloration uniform or darkened dorsally. 


a 
ETYMOLOGY: From the Greek 8 PX US 
(brachys), short, TW MA (soma), body, 
and op cf (ophis; masculine), serpent. 


DISTRIBUTION: Known from four central and 
western Pacific and a single eastern Atlantic spe- 
cies. Gunther’s (1870) report of B. crocodilinus 
from the Galapagos Islands was probably er- 
roneus (Rosenblatt et al., 1972). 


REMARKS: The species of Brachysomophis are 
clearly in need of revision (Schultz, 1943; Smith, 
1962) but | am lacking the material to make the 
required comparisons. Brachysomophis atlanticus 
Blache and Saldana (1972) was recently described 
from Senegal. The Pacific species of Brachysomo- 
phis include: Ophisurus cirrhocheilos Bleeker*, 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


O. corcodilinus Bennett*, Brachysomophis hen- 
shawi Jordan and Snyder, and B. sauropsis 
Schultz. The last three nominal species are quite 
similar and perhaps conspecific. B. cirrhocheilos 
differs from those species in apparently lacking 
the postorbital depression, in its longer pectoral 
fin, and faintly banded coloration (fide Deraniya- 
gala, 1929). The neurocranium and suspensorium 
of B. henshawi were illustrated by Gosline 
(1951a: fig. 2). 

The formation of the fused postorbital strut in 
species of Brachysomophis is functionally con- 
vergent with that of Ophichthus zophochir, how- 
ever the third, dorsal-most postorbital of B. 
sauropsis is longer than the second, and enlarged 
anteriorly to form a cup that braces the eye. 


Cirrhimuraena Kaup 


Cirrhimuraena Kaup 1856; 51 (27). (Type species; 
C. chinensis Kaup 1856, by monotypy.) 

Jenkinsiella Jordan and Evermann 1905: 83. (Type 
species; Microdonophis macgregori Jenkins 
1903, by original designation.) 

Calamuraena Whitley 1944: 261. (Type species; 
Ophichthys calamus Gunther 1870, by original 
designation.) 

Calamuraena Gunther 1870, sic Rosenblatt and 
McCosker 1970: 496, lapsus pro Calamuraena 
Whitley 1944. 


DESCRIPTION: General characteristics those of 
Ophichthus. Differences include: DFO generally 
on head or above GO; pectoral fin moderately 
to well developed; upper lip with numerous 
cirri; pop® generally present; teeth conical, 
pointed but not enlarged, often multiserial in 
jaws and on vomer, those of intermaxillary and 
vomer discontinuous; skull subtruncate posteri- 
orly (more so than in Ophichthus); PO strut ab- 
sent; SO and frontal crests weakly developed; 
maxilla elongate posteriorly; otic bulla not 
strongly developed; actinosts absent, or 1-2; col- 
oration uniform, often darkened dorsally. 


ETYMOLOGY: From the Latin cirrus (cirrh is a 
wrong but very common form of cirr, mistakenly 
considered to be from the Greek 4e PEOS 
[kirrhos] meaning yellow or tawny), tendril, and 
Muraena, a genus of muraenid eels. 


DISTRIBUTION: An Indo-Pacific and Red Sea 
genus, with ca. 9 valid species. 


REMARKS: Cirrhimuraena is broadly defined to 
include species that have been referred to Cala- 
muraena and Jenkinsiella. Subgeneric lines within 


75 


Cirrhimuraena may be separated on an external 
morphological basis in the following manner: 


Mandibular and vomerine teeth uniserial; 
pop® absent (not determined for all species) 
Baca Be sci it 3 ee ee subgenus Jenkinsiella 


Mandibular and vomerine teeth multiserial; 
pop® present (not determined for all species) 
Re iS Rie ea eee S83 subgenus Cirrhimuraena 


The subgenus Jenkinsiella Jordan and Evermann 
includes Microdonophis macgregori Jenkins, 
Ophichthys playfairii Gunther*, and may include 
Jenkinsiella oliveri Seale* and J. inhacae Smith*. 
The remaining species belong in the subgenus 
Cirrhimuraena Kaup (which includes Calamur- 
aena Whitley), including: Cirrhimuraena chinen- 
sis Kaup, C. taeniopterus Bleeker, C. paucidens 
Herre and Myers (=C. chinensis?), Ophichthys 
calamus Gunther, and Ophisurus cheilopogon 
Bleeker*. Cirrhimuraena may merit further gen- 
eric division, particularly if confirmed by an 
osteological study of all the included species. 

Nelson’s (1966a: 395) description of the fifth 
ceratobranchial of C. macgregori as having ‘‘the 
proximal portion of Cs cartilaginous... (and) the 
distal portion extending posteriorly as a thin 
filament of cartilage’’ does not agree with my 
findings on available specimens. In them the 
distal portion is well ossified. Gosline (1951a: 
fig. 6) has illustrated the neurocranium and sus- 
pensorium of C. macgregori. 


Echelus Rafinesque 


Echelus Rafinesque 1810a: 63. (Type species; E. 
punctatus Rafinesque 1810a = Muraena myrus 
Linnaeus 1758, by Bleeker 1864 as first reviser.) 

Myrus Kaup 1856: 53 (31). (Type species; M. vul- 
garis Kaup 1856 = Muraena myrus Linnaeus 
1758, by monotypy.) 


DESCRIPTION: General characteristics those of 
Ophichthus. Differences include: DFO before 
pectoral tips; median fins continuous around 
tail; head pores reduced, pop® present, temporal, 
postorbital, and interorbital pores absent; ceph- 
alic surface sensory pores well developed; teeth 
multiserial, small, nearly granular, intermaxillary 
teeth continuous with those of vomer; PO strut 
absent, but a cartilaginous antorbital strut present 
in E. myrus; SO and frontal crests moderately 
developed; maxilla slender posteriorly; branch- 
iostegal rays few; coloration nearly uniform, 
darker dorsally. 


76 


2, 
ETYMOLOGY: From the Greek éypenue 
(enchelys, considered either feminine or mascu- 
line, regarded as masculine by Rafinesque, Kaup, 
and Bleeker), eel. 


DISTRIBUTION: Known from a shallow and a 
deep-water species from the Mediterranean and 
eastern Atlantic, respectively, E. myrus (Linnaeus) 
and Myrus pachyrhynchus Vaillant. 


REMARKS: The osteology of the species of 
Echelus indicates that they are closely related to 
species of Ophichthus, with particular similarities 
evidenced in the hyoid arches, gill arches, and 
neurocrania. The conspicuous presence of a 
caudal fin led previous workers to consider 
Echelus to belong to a family or subfamily sep- 
arate from Ophichthus. The tail fin is clearly a 
superficial condition that does not merit such 
suprageneric separation, and thus the family 
name Echelidae is referred to the synonymy of 
the Ophichthidae, and the subfamily Echelinae 
to that of the Ophichthinae. 

The species of Echelus are easily separable 
from other ophichthins on the basis of two char- 
acters, the presence of extremely visible caudal 
fin rays, and the head pore reduction. The pro- 
duced caudal fin appears to be a primitive re- 
tention of an ancestral conditon, although the 
hard-pointed tail tip characteristic of other oph- 
ichthines was probably achieved early in the 
evolution of the subfamily. The absence of the 
temporal, postorbital, and interorbital pores (as 
noted by Gosline, 1952) probably reflects the 
loss of these openings, in that the cephalic later- 
alis canals and frontal commissures of E£. myrus 
(Allis, 1903: figs. 5-8, and personal observation) 
and £. pachyrhynchus (personal observation) 
have been retained. The antorbital cartilaginous 
strut of E. myrus, as identifed by Gosline (1952), 
is not present in E. pachyrhynchus and does not 
appear ot be an important indicator of phylo- 
geny. Gosline (1952: 133) suggested that “Ech- 
elus myrus appears, despite specializations, to be 
by far the most primitive (or generalized) ophich- 
thid known. In fact, to a considerable degree it 
fills in the gap between the Ophichthidae and 
the Congridae.” | concur with Gosline in consid- 
ering Echelus to be a very generalized (primitive) 
ophichthid, however its gill arch reduction, labial 
posterior nostril, and hyoid apparatus suggest 
that it is well separated from any congrid-like 
ancestor. 

The neurocranium of Echelus myrus has been 
discussed and illustrated by Storms (1896, as 


SERIES 4, V. 41, #1 McCOSKER — EELS 


Myrus vulgaris) and by Gosline (1952), and the 
external morphology of £. myrus and E. pachy- 
rhynchus by Blache (1968). 


Echiophis Kaup 


Echiophis Kaup 1856a: 46. (Type species; Ophis- 
urus intertinctus Richardson 1844b, by mono- 
typy.) 

Echiopsis Kaup 1856b: 13. Emend. pro Echiophis 
Kaup 1856a. 

Crotalopsis Kaup 1860: 12. Also spelled Crotalo- 
pis, Crotalophis, by other authors. (Type spe- 
cies; C. punctifer Kaup 1860, by monotypy.) 

Macrodonophis Poey 1867: 251. (Type species; 
Conger mordax Poey 1861, by monotypy.) 


DESCRIPTION: Body cylindrical, moderately 
elongate, laterally compressed posteriorly; body 
shorter than tail; snout short, subconical, slightly 
constricted near tip; jaws elongate and sub- 
equal; eye moderate; posterior nostril in a short 
tube before and beneath eye; GO vertical, lateral 
and elongate, = ishtmus; DFO behind pectoral 
tips; tail tip blunt; pop*® present; LL ossicles sep- 
arated at pores; teeth strong, pointed and bi- 
serial, largest anteriorly in jaws and on ethmoid, 
those of intermaxillary and vomer biserial and 
nearly continuous; skull subtruncate posteriorly, 
preorbital region reduced, orbit large; PO fused 
to skull and maxilla, forming a strut; nasals 
slender, nasal cartilage reduced or absent; SOC 
absent; maxilla elongate, rounded posteriorly, 
toothed along most of its length; opercular mar- 
gins entire, subopercle scythe-shaped; suspen- 
sorium posteriorly inclined, jaw angle ca. 85°; 
hyomandibular ridged and elongate posterodor- 
sally for muscle attachment; otic bulla well de- 
veloped; PG broad posteriorly, free and terete 
anteriorly; hyoid arch slender, HH separated 
from CH by a narrow gap; branchiostegal rays 
numerous, all slender, unbranched, and along 
hyoid; UH a spike posteriorly; C; ossified, UPs- 
UP, fused; pectoral girdle and fin developed, Sc 
and Co large, actinosts present; caudal vertebrae 
slightly more than precaudal; coloration gener- 
ally strongly spotted. 


aif 
ETYMOLOGY: From the Greek ees 
(echis), viper, and O PL (ophis; mascu- 
line), serpent. Kaup emended the suffix to ¥ 
tf (Opsis), appearance. 


DISTRIBUTION: A largely New World genus con- 
taining 3-5 Atlantic and Pacific species. Included 
are: Ophisurus intertinctus Richardson (WA, and 


_Cyclophichthys Whitley 1951: 


— 


- apparently 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


EA as Mystriophis cruetzbergi Cadenat), Conger 
mordax Poey (WA), Crotalopsis punctifer Kaup 
(WA), and an undescribed eastern Pacific species 
ranging from the Northern Gulf of California to 
Panama. Incertae sedis: Mystriophis blastorhinos 
Kanazawa (WA). 


REMARKS: The species of Echiophis form a dis- 
tinctive complex within the Ophichthini, and are 
difficult to separate at the specific level. The At- 
lantic species are separable from each other on 
the basis of spot size, yet a continuous grade is 
clearly present. Opinions as to the distinctiveness 
of the Atlantic species have been presented (Jor- 
dan and Davis, 1891; Springer and Allen, 1932; 
Ginsburg, 1951) yet a conclusive study is still 
lacking. The morphology of eastern Atlantic 
specimens of EF. intertinctus is described by 
Blache (1971). 


Elapsopis Kaup 


Elapsopis Kaup 1856: 45 (9). Emended to Elaps- 
opsis by other authors. (Type species; Ophi- 
surus versicolor Richardson 1844, by mono- 
typy.) 

392. Described as 

a subgenus of Malvoliophis Whitley 1934. 

(Type species; Ophichthus cyclorhinus Fraser- 

Brunner 1934, by original designation.) 


DESCRIPTION: General characteristics those of 
Leiuranus. Differences include: anterior nostril 
in a short tube, its rim flared distally; DFO above 
or slightly before GO; pectoral fin reduced, ~ 
GO length; vomerine teeth present; opercular 
margin fringed; C; ossified; actinosts of pectoral 
girdle present. 


ETYMOLOGY: From the Latin e/aps (masculine), 
a snake, and opis, an erroneous spelling of the 
Greek © roy (opsis), Meaning appearance. 


DISTRIBUTION: Included are two, probably con- 
specific, species, from Australia, Lord Howe Is- 
land, and the southern Caroline Archipelago. 


REMARKS: This genus is provisionally recognized 
as distinct from Leiuranus on the basis of the 
secondary ossification of the fifth 
ceratobranchial and the presence of actinosts. 


_ The occasional presence of 1-2 vomerine teeth 
in specimens of L. semicinctus approaches the 


| 


condition of certain specimens of EF. versicolor 
and E. cyclorhinus. Further investigation may re- 
sult in the recognition of Elapsopis as a subgenus 
of Leiuranus. 


Te, 


Evips McCosker 


Evips McCosker 1972: 113. (Type species; E. per- 
cinctus McCosker 1972, by original designa- 
tion.) 


DESCRIPTION: Body moderately elongate, cyl- 
indrical, laterally compressed posteriorly; body 
slightly longer than tail; snout blunt, conspicu- 
ously papillate; lower jaw included; eye large; 
DFO behind GO; pectoral fin minute; tail tip 
pointed; pop® absent; LL ossicles continuous, but 
heavily fractionated; teeth pointed, uniserial, ex- 
cept those of maxilla which are biserial, largest 
at intermaxillary which are continuous with those 
of vomer; skull subtruncate posteriorly, orbit 
large; SOC pointed posteriorly; maxilla moder- 
ately produced, slender posteriorly; suspensor- 
ium nearly vertical, jaw angle ca. 95°; hyoid 
stout, HH separated from CH by a gap; branchi- 
ostegal rays numerous, all slender and associated 
with hyoid; UH a spike posteriorly; C; present, 
UP3-UP4 separate; pectoral girdle reduced to a 
slender Cl (SCI not visible in radiograph), actin- 
osts absent; caudal = precaudal vertebrae; col- 
oration strong banded. 


ETYMOLOGY: From the Greek € % (eu, 
latinized to ev for euphony before a vowel), 
good, and Ga af (ips; masculine), a worm. 


DISTRIBUTION: A_ single species known only 
from the type specimen from the Southern Car- 
oline Archipelago. 


Leiuranus Bleeker 


Leiuranus Bleeker 1853a: 24. (Type species; L. 
lacepedii Bleeker 1853a = Ophisurus semi- 
cinctus Lay and Bennett, by monotypy.) 

Stethopterus Bleeker 1853a: 24. (Type species; 
Ophisurus (Sphagebranchus) vimineus  Rich- 
ardson 184442 = Ophisurus semicinctus Lay 
and Bennett 1839, by monotypy.) 

Machaerenchelys Fowler 1937: 85. (Type species; 
M. vanderbilti Fowler 1937 = Ophisurus semi- 
cinctus Lay and Bennett 1839, by original des- 
ignation.) 

DESCRIPTION: Body moderately elongate, cy- 

lindrical, laterally compressed posteriorly; body 

and tail subequal; snout conical, flattened and 
grooved on underside; lower jaw included; eye 
moderate; anterior nostril in a short tube, pos- 
terior along lip margin; GO vertical, crescentic, 
shorter than isthmus; median fins low, expanded 
before tail tip; DFO above GO; pectoral fin 
moderately developed; tail tip sharply pointed; 


78 


pop® absent; LL ossicles narrowly separated at 
pores; teeth small, recurved, uniserial, absent 
from vomer or 1-2 small teeth; neurocranium 
subtruncate posteriorly, orbit large; PO strut ab- 
sent; nasals and nasal cartilage moderately de- 
veloped; frontal-parietal ridge and SOC devel- 
oped; maxilla short, articulated ca. mid-vomer; 
maxillae closely apposed anteromedially (but not 
abutting as per Gosline, 1951a: 301); suspensor- 
ium anteriorly inclined, jaw angle ca. 100°; otic 
bulla well developed; PG slender, braced pos- 
teriorly by hyomandibular, pointed anteriorly; 
HH separated from CH by a short gap; branchio- 
stegal rays numerous, slender (distal-most broad- 
ened slightly), unbranched, and associated with 
hyoid (except for innermost rays); UH split an- 
teriorly (less so than in Phyllophichthus), pro- 
duced posteriorly as a spike; gill arches weak, 
Cs absent, anterior half of H: ossified; pectoral 
girdle reduced, actinosts absent; precaudal ver- 
tebrae more numeyous than caudal; coloration 
strongly banded. 


rs 
ETYMOLOGY: From the Greek AECL (leios), 
smooth, and 0 UP KUS, (ouranos; masculine), 
sky, in reference to the toothless vomer. 


DISTRIBUTION: A single species, widespread in 
the central and western Pacific and Indian 
oceans. 


REMARKS: Gosline (1951a: fig. 4) has illustrated 
the neurocranium and suspensorium of L. semi- 
cinctus. 


Malvoliophis Whitley 


Malvoliophis Whitley 1934: 154. (Type species; 
Bascanichthys hemizona Ogilby = Ophichthys 
pinguis Gunther 1872, by original designation.) 


DESCRIPTION: Body elongate, subcylindrical, 
laterally compressed; body shorter than tail; 
snout developed, subconical, a short groove on 
underside; lower jaw included; eye large; an- 
terior nostril tubular; DFO before GO; pectoral 
fin moderately developed; pop* absent; LL os- 
sicles widely separated at pores; teeth conical, 
not sharply pointed, unserial in jaws, largest at 
intermaxillary which are widely separated from 
those of the vomer; skull subtruncate posteriorly, 
orbit large; PO strut absent; maxilla elongate, 
slender posteriorly; opercular series weakly de- 
veloped; suspensorium nearly vertical, jaw angle 
ca. 95°; otic bulla moderately developed; PG 
broad _ posteriorly, slender anteriorly; hyoid 
slender, HH separated from CH by a suture; 


SERIES 4, V.41,#1 McCOSKER — EELS 


branchiostegal rays numerous, slender and 
slightly flattened, the anteriormost ray slightly 
enlarged; UH broad anteriorly, a short spike 
posteriorly; Cs; weakly ossified (cartilaginous?), 
UP;-UPs fused; pectoral girdle reduced, lacking 
Sc, Co, and actinosts; caudal vertebrae more 
numerous than precaudal; coloration weakly 
banded and spotted. 


ETYMOLOGY: From Malvolio, Lady  Olivia’s 
steward in Shakespeare’s Twelfth Night, and 

opts (ophis; masculine), serpent. The 
banded coloration of M. pinguis, the generic 
type, suggests the cross-gartered legs and yellow 
socks worn by Malvolio (G. P. Whitley, personal 
communication). 


DISTRIBUTION: A single species, reported from 
southern Queensland, New South Wales, and 
Lord Howe Island. 


Myrichthys Girard 


Myrichthys Girard 1859: 58. (Type species; M. 
tigrinus Girard 1859, by monotypy.) 

Chlevastes Jordan and Snyder 1901: 867. (Type 
species; Muraena colubrina Boddaert 1781, by 
original designation.) 


DESCRIPTION: General characteristics those of 
Pisodonophis and Ophichthus. Differences  in- 
clude: snout short (for an ophichthin), broad 
from above; DFO well before GO; pectoral fin 
short, broad-based (fig 34); pop* absent; teeth 
molariform or granular, multiserial on jaws and 
vomer, largest at intermaxillary, which are nar- 
rowly separated from those of vomer; PO strut 
absent; maxilla elongate, slender posteriorly; 
teeth of pharyngeal plates generally small and 
pavement like; pectoral girdle reduced, Co re- 
duced, Sc and actinosts absent; coloration either 
spotted, banded, or both. 


ETYMOLOGY: From the Greek paupPos 
(myrus), eel, and Ge (2) ba 6 (ichthys; 


masculine), fish. 


DISTRIBUTION: A circumtropical genus currently 
including seven valid species. The genus may be 
expanded to 12 should certain insular popula- 
tions of M. maculosus and M. xystrurus prove to 
be distinct. Included in Myrichthys are: M. 
bleekeri Gosline (a substitute name for Ophi- 
surus fasciatus var. semicinctus Bleeker, preoc- 
cupied) (IP), M. sp. (EP), Muraena acuminata 
Gronow (WA), M. colubrina Boddaert (IP), M. 
maculosa Cuvier (IP), Pisodonophis oculatus Kaup 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


(WA), Ophisurus pardalis Valenciennes (EA)*, 
and Ophichthys xystrurus Jordan and Gilbert (EP) 
(see Remarks). 


REMARKS: Muraena tigrina Ruppell (1826), de- 
scribed from the Red Sea, is currently placed in 
the synonymy of Myrichthys maculosus (Cuvier). 
This creates a homonymy with the eastern Pacific 
Myrichthys tigrinus Girard (1859), the type spe- 
cies of the genus. Myrichthys xystrurus (Jordan 
and Gilbert, 1882) is the next available name for 
the eastern Pacific species. 

The genera Pisodonophis and Myrichthys are 
offshoots from a generalized Ophichthus-like an- 
cestor. Within the examined species of Pisodon- 
ophis a trend in characters is evident, with P. 
cancrivorus the most Ophichthus-like and P. 
daspilotus the most Myrichthys-like (the char- 
acters however, are discontinuous enough to al- 
low generic separation). These trends include: 
the transformation from pointed to molariform 
dentition; the advancement of the DFO; the re- 
duction in length and basal broadening of the 
pectoral fin; and the reduction of the coracoid 
and loss of the scapula. 

Harry (1949) has pointed out that the type 
locality of M. tigrinus is properly Adair Bay, Mex- 
ico, not Oregon. 

Gosline (1951a) has illustrated the neurocran- 
ium, vertebrae, gill arches, and pectoral girdle 
of M. maculosus. 


Mystriophis Kaup 


Mystriophis Kaup 1856: 45 (10). (Type species; 
Ophisurus_ rostellatus Richardson 1844, by 
monotypy.) ; 


DESCRIPTION: Body cylindrical, moderately 
elongate, laterally compressed posteriorly; body 
shorter than tail; snout short, subconical, con- 
stricted near tip; jaws elongate, the lower slightly 
inferior; eye moderate; posterior nostril at edge 
of upper lip, covered by a flap; GO vertical, lat- 
eral and elongate, = isthmus; DFO above or be- 
hind pectoral tips; tail tip blunt; pop*® absent, 
secondary cephalic pores well developed; LL 
ossicles separated at pores; teeth strong, pointed, 
biserial, largest anteriorly in jaws and on eth- 
moid, intermaxillary and uniserial vomerine teeth 
nearly continuous; coloration nearly uniform, 
darkened dorsally. 


ETYMOLOGY: From the Greek pau oT Ploy 
(mystrion), a spoon, and Spe (ophis; 
masculine), serpent, in relation to the snout 
shape of M. rostellatus, the generic type. 


7/2) 


DISTRIBUTION: Two eastern Atlantic and one 
Japanese species are provisionally placed in My- 
striophis (see following Remarks). 


REMARKS: Specimens of Mystriophis were un- 
available for osteological examination, therefore 
the above description was prepared from Blache 
(1971). The genera Mystriophis and Echiophis 
have been separated by recent authors (Gins- 
burg, 1951; Kanazawa, 1963; Rosenblatt and 
McCosker, 1970; Blache, 1971) on the basis of 
the snout shape and vomerine dentition. M. ros- 
tellatus Richardson and M. crosnieri Blache, from 
the eastern Atlantic, are undoubtedly congeneric 
and appear to be recently derived from an 
Ophichthus-like ancestor (evidenced in the pop? 
absence, secondary cephalic pore development, 
moderate snout length, and body coloration). 
The species of Echiophis however, appear to 
have been derived separately from a rather dif- 
ferent Ophichthus-like ancestor (pop* present, 
secondary cephalic pores absent, shorter snout, 
and spotted coloration). Ophisurus porphyreus 
Temminck and Schlegel, from Japan, tentatively 
placed in Brachysomophis by Kanazawa (1963), 
probably belongs in Mystriophis. M. blastorhinos 
Kanazawa, from French Guiana, is either an 
Echiophis or belongs in a separate genus. Kana- 
zawa’s illustration of the pore pattern of M. blas- 
torhinos is somewhat inaccurate; pop*® and a 
single, rather than a double, interorbital pore are 
present (Kanazawa, in litt.). 


Ophichthus Ahl 


Ophichthus Ahl 1789: 5. Emended to Ophichthys 
by other authors. (Type species; Muraena 
ophis Linnaeus 1758, by original designation.) 

Innominado Parra 1781: 96. A junior synonym of 
Muraena ophis Linnaeus 1758, non-binomial. 

Ophis Turton 1807: 87. (Type species; ““O. macu- 
lata . . . Spotted Serpent. Shaw Zool., iv. p. 22 
. . . Bloch t. 154,” presumably based on Mur- 
aena ophis Linnaeus 1758, by monotypy.) 

Cogrus Rafinesque 1810a: 62. (Type species; C. 
maculatus Rafinesque 1810a, by monotypy.) 

Ophithorax McClelland 1844: 212. (Type spe- 
cies; Ophisurus ophis Lacépede 1800, presum- 
ably based on Muraena ophis Linnaeus 1758, 
by Jordan, 1919b, as first reviser.) 

Centrurophis Kaup 1856: 42 (2). (Type species; 
Ophisurus spadiceus Richardson 1844, as a 
misidentification of Ophichthys cephalazona 
Bleeker 1864, by Jordan, 1919b, as first re- 
viser.) 

Poecilocephalus Kaup 1856. 43 (5): (Type spe- 


80 


cies; P. bonaparti Kaup 1856, by monotypy.) 

Microdonophis Kaup 1856: 43 (6). (Type species; 
M. altipinnis Kaup 1856, by monotypy.) 

Coecilophis Kaup 1856: 44 (6) (Type species; 
Ophisurus compar Richardson 1844 = Ophi- 
surus apicalis Bennett 1830, by monotypy.) 

Herpetoichthys Kaup 1856: 44 (7). (Type species; 
H. ornatissimus Kaup 1856, by Jordan, 1919b, 
as first reviser.) . 

Muraenopsis LeSueur, sic Kaup 1856: 46 (11), 
lapsus pro Muraenophis Lacépéde 1825. 

Scytalophis Kaup 1856: 46 (13). (Type species; S. 
magnioculis Kaup 1856, by Jordan, 1919b, as 
first reviser.) 

Leptorhinophis Kaup 1856: 46 (14). (Type spe- 
cies; Ophisurus gomesi Castelnau 1855, by 
Jordan, 1919b, as first reviser.) 

Cryptopterus Kaup 1860: 11. (Type species; C. 
puncticeps Kaup 1860, by monotypy.) 

Uranichthys Poey 1867: 256. (Type species; Mur- 
aena hauannensis Bloch and Schneider 1801 = 
Muraena ophis Linnaeus 1758, by Jordan and 
Davis, 1891, as first revisers.) 

Oxydontichthys Poey 1880: 254. (Type species; 
Ophichthys macrurus Poey 1880 = Ophisurus 
gomesi Castelnau 1855, by original designa- 
tion.) 

Omochelys Fowler 1918: 3. Described as a sub- 
genus of Pisodonophis Kaup. (Type species; 
Pisodonophis cruentifer Goode and Bean 1895, 
by original designation.) 

Syletor Jordan 1919a: 343. (Type species; Piso- 
odonophis cruentifer Goode and Bean 1895, 
by original designation.) 

Styletor Jordan 1919a, sic Jordan, Evermann, and 
Clark 1930: 86, lapsus pro Syletor Jordan 
1919a. 

Acanthenchelys Norman 1922: 296. (Type spe- 
cies; A. spinicauda Norman 1922, by original 
designation.) 

Cryptopterenchelys Fowler 1925: 1. Described 
as a subgenus of Ophichthus Ahl. (Type spe- 
cies; Cryptopterus puncticeps Kaup, as a sub- 
stitute name for Cryptopterus Kaup 1860, pre- 
occupied.) 

Zonophichthus Whitley 1930: 250. (Type spe- 
cies; Ophichthys cephalazona Bleeker 1864, 
by original designation.) 

Gisenchelys Fowler 1944: 188. Described as a 
subgenus of Ophichthus Ahl. (Type species; 
Ophichthys zophochir Jordan and_ Gilbert 
1881, by original designation.) 

Syletophis Whitley 1950: 44. Substitute name 
for Syletor Jordan 1919a, preoccupied. 


SERIES 4, V. 41,#1 McCOSKER — EELS 


Antobrantia Ypiranga Pinto 1970: 13. (Type spe- 
cies; A. ribeiroi Ypiranga Pinto 1970 = Mur- 
aena ophis Linnaeus 1758, by original designa- 
tion.) 


DESCRIPTION: Body moderately to very elongate, 
laterally compressed posteriorly; body shorter 
than tail; snout moderately developed, subconi- 
cal; lower jaw included; eye moderate to large; 
anterior nostril tubular, posterior along lower 
edge of lip or opening into mouth; DFO above 
or behind GO, but generally before pectoral 
tips; pectoral fin well developed; LL ossicles sep- 
arated at pores; dentition variable, from num- 
erous, small, and multiserial to few, large, and 
uniserial or biserial, never caniniform; teeth larg- 
est at intermaxillary and on vomer and some- 
times separated by a short gap; skull subtruncate 
posteriorly (fig. 2), orbit large; PO strut devel- 
oped in some species (fig. 14); nasals and nasal 
cartilage moderately developed; frontal and SO 
crests moderately to well developed; maxilla 
elongate, slender or rounded posteriorly (figs. 14, 
16); opercular series well developed, their mar- 
gins entire (fig. 14); suspensorium nearly vertical 
to posteriorly inclined; otic bulla well devel- 
oped; PG broad posteriorly, free and terete an- 
teriorly; hyoid slender, HH separated from CH 
by a narrow gap (fig. 17); branchiostegal rays 
slender, often numerous, unbranched and along 
arch; UH a spike posteriorly; gill arches com- 
plete, C; present, UP3-UP.s separate (fig. 18); pec- 
toral girdle well developed, actinosts present 
(fig. 19A); caudal vertebrae more numerous than 
precaudal; coloration generally uniform, although 
banded or spotted species may be included. 


Xf 
ETYMOLOGY: From the Greek O MLS 
(ophis), snake, and ix evs (ichthus, more 
correctly written ichthys; masculine), fish. 


DISTRIBUTION: A circumtropical genus with ap- 
proximately 55 species. 


REMARKS: Ophichthus contains approximately 
55 valid species, for which no fewer than 25 
nominal genera have been erected. The genera 
Gisenchelys Fowler and Zonophichthus Whitley 
are here included in the synonymy of Ophich- 
thus. Hubbs’ (1932) inclusion of Ophis Turton 
in the synonymy of Ophichthus was neither in- 
cluded nor commented upon by Bohlke and 
Robins (1959) in their synonymy of Ophichthus 
ophis. Bohlke and Menezes (in litt.) have found 
Antobrantia, type species A. ribeiroi Ypiranga 
Pinto, to be an exact synonym of Ophichthus 
ophis. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


The generic concepts of earlier authors, par- 
ticularly Kaup, are nearly all based on differ- 
ences in coloration or dentition. | have at- 
tempted to examine as many of these subgenera 
as possible. As broadly defined herein, Ophich- 
thus exhibits a wide variety of morphological 
conditions, yet among the diverse types ex- 
amined, | have found no accompanying differ- 
ences that are clearly generic. It is particularly 
perplexing however, that the few osteological 
variations within the genus that might merit gen- 
eric recognition are present in O. triserialis, O. 
zophochir, and the type species, O. ophis. These 
concern the development of the postorbital strut 
and the posterior shortening of the maxilla. 

| have attempted to identify subgeneric line- 
ages within Ophichthus, using available generic 
synonyms as subgeneric names. The following 
analysis however, must be regarded as strictly 
provisional: 


Body = tail; DFO above or before GO; 
teeth minute and uniserial throughout; col- 
oration of several species strongly spotted)... 
nonseee subgenus Microdonophis Kaup 


Body shorter than tail; DFO behind GO, 
generally above pectoral tips; teeth larger, 
ier GANS) ee eee 


Anterior nostril a broad tube, flared at 
tip; intermaxillary block hooked down- 
ward, and intermaxillary teeth directed 
horizontally backward; tail tip hard and 
sharply-pointed; median fins conspicu- 
ously expanded before tail tip -................... 
a subgenus Centrurophis Kaup 


Anterior nostril tubular, not flared distally; 
intermaxillary teeth erect, with tips hook- 
ed back; tail tip hard and blunt, not 
sharply-pointed; median fins conspicu- 
spicuously expanded before tail tip -....... 
ah ee ee subgenus Centrurophis Kaup 


Postorbital bones tightly sutured and 
bracing maxilla, forming a strut; maxilla 
rounded posteriorly; jaw and vomerine 
dentition bi- or triserial -....................... 
ie eee ee A subgenus Ophichthus Ahl 


Postorbital bones not tightly sutured 
and not forming a strut with maxilla; 
maxilla elongate and slender posteri- 
orly; dentition variable, either uniserial 
OTN UNI CUS tile lee Se eee es 
oe een subgenus Coecilophis Kaup 


Included in the subgenus Microdonophis are: 
M. altipinnis Kaup (IP), M. erabo Jordan and 


81 


Snyder (IP), Ophichthys polyophthalmus Bleeker 
(IP)*, and O. melanochir Bleeker (IP). 

Included in the subgenus Centrurophis are the 
generic types of Zonophichthus and_ probably 
Poecilocephalus. \Included species: Ophichthys 
cephalazona Bleeker (IP), and Poecilocephalus 
bonaparti Kaup (IP)*. 

Included in the subgenus Ophichthus are the 
generic types of Innominado, Ophis, Cogrus, 
Uranichthys, Ophithorax, and Gisenchelys. In- 
cluded species: Muraena ophis Linnaeus (WA, 
EA), Muraenopsis triserialis Kaup (EP), and Oph- 
ichthys zophochir Jordan and Gilbert (EP). A 
complete synonymy of O. ophis is provided by 
Bohlke and Robins (1959). 

Included in the subgenus Coecilophis is Ophi- 
surus apicalis Bennett (IP), and presumably many 
other species listed below as incertae sedis. 

Incertae sedis: Acanthenchelys spinicauda Re- 
gan (WA)*, Cogrus maculatus Rafinesque (M)*, 
Conger urolophus Schlegel (IP), Cryptopterus 
puncticeps Kaup (WA), Echelus rufus Rafinesque 
(M)*, Omochelys marginatus Fowler (WA) (= 
Ophichthus cruentifer 2), Ophichthus asakusae 
Jordan and Snyder (IP), O. manilensis Herre (IP)*, 
O. melanoporus Kanazawa (WA), O. multiserialis 
Norman (IP)*, O. retifer Fowler (IP)*, O. roseus 
Tanaka (IP)*, O. stenopterus Cope (IP)*, Ophich- 
thys ascensionis Studer (WA)*, O. ater Peters (EP)*, 
O. callensis Gunther (EP), O. derbeyensis Whitley 
(IP)*, O. episcopus Macleay (IP)*, O. evermanni 
Jordan and Snyder (IP), O. frontalis Garman (EP), 
O. garretti Gunther (IP)*, O. limkouensis Chen 
(IP)*, O. macrops Gunther (IP)*, O. madagascari- 
ensis Fourmanoir (IP)*, O. melanochir Bleeker (IP), 
O. pacifici Gunther (EP), O. retropinnis Eigen- 
mann (WA)*, O. unicolor Regan (IP)*, O. woo- 
suitingi Chen (IP)*, Ophisurus celebicus Bleeker 
(IP)*, O. gomesii Castelnau (WA), O. grandoculis 
Cantor (IP)*, O. macrochir Bleeker (IP), O. mar- 
ginatus Peters (IP)*, O. parilis Richardson (WA)*, 
O. reguis Richardson (EA)*, O. remiger Valen- 
ciennes (probably EP, no type locality given)*, 
O. rutidoderma Bleeker (emended by most au- 
thors to rhytioderma), (IP)*, O. rutidodermatoides 
Bleeker (emended by most authors to rhytioder- 
matoides), (IP), Pisodonophis cruentifer Goode 
and Bean (WA), Scytalophis magnioculis Kaup 
(WA)*, and two undescribed Eastern Pacific spe- 
cies. 


Ophisurus Lacépede 


Ophisurus Lacépéde 1800: 195. (Type species; 
Muraena serpens Linnaeus 1758, as restricted 


82 


by Risso 1826.) 

Leptognathus Swainson 1838: 334. (Type species; 
L. oxyrhynchus Swainson 1838 = Muraena ser- 
pens Linnaeus 1758, by original designation.) 

Leptorhynchus Smith 1847: 244. (Type species; 
L. capensis Smith 1847 = Muraena serpens 
Linnaeus 1758.) 

Anepistomon Gistel 1848: ix. A substitute name 
for Leptorhynchus Smith 1847, preoccupied. 


DESCRIPTION: General characteristics those of 
Ophichthus. Differences include: body very 
elongate; snout attenuate, jaws elongate, slender, 
incapable of closing completely in adults; lower 
jaw included; anterior nostril lateral, at mid- 
snout (fig. 24B), posterior on outer lip and cov- 
ered by a flap; DFO beyond tips of pectoral fins; 
cephalic surface sensory papillae well developed 
(figs. 24B, C); teeth conical, nearly uniserial, en- 
larged along vomer; maxilla elongate, slender 
and produced posteriorly; coloration uniform, 
darker dorsally. 


ETYMOLOGY: From the Greek & pey (ophis), 
serpent, and OY PX (oura), tail, and -of 
(-us), masculine suffix. 


DISTRIBUTION: Presumably a single species, 
known from the Mediterranean, eastern Atlantic, 
Cape of Good Hope, the western and central 
Indian Ocean, eastern Australia, New Zealand, 
and Japan. Ophisurus macrorhynchus Bleeker 
1852, from Japan, differs slightly in morphometry 
from the Mediterranean Muraena serpens, yet a 
conclusive comparison has not been made. 


REMARKS: The Atlantic and Pacific populations 
of O. serpens, if indeed conspecific, represent 
the most disjunct of known ophichthid distribu- 
tions. Historically, the Mediterranean population 
could be explained as a relict of a Tethyan spe- 
cies. Parallels may be found in the disjunct pop- 
ulations of other eel species (see D. Smith, 1969), 
such as Kaupichthys diodontus, which presum- 
ably ranges from the western Atlantic, and the 
central and western Pacific, and Indian oceans 
J. L. B. Smith, 1965), and Dysomma anguillare, 
presumably ranging from the tropical western 
Atlantic, Indian Ocean, and off Japan and China 
(Robins and Robins, 1970). 

The identity of Ophichthus multiserialis Nor- 
man 1939 from the Gulf of Aden, included by 
J. L. B. Smith (1962: 455) in Ophisurus, has not 
been determined. 

Oxystomus Rafinesque (1810b: 62) has been 
considered by most authors to be synonymous 
with Ophisurus. Castle (1969) however, has con- 


SERIES 4, V.41, #1 McCOSKER — EELS 


firmed Bertin’s (1935) suggestion that the type 
species, Oxystomus hyalinus, is based on a post- 
metamorphic Serrivomer rather than a_ larval 
Ophisurus serpens. 

The external morphology and osteology of 
Ophisurus serpens (as O. novaezelandiae Hector) 
was described by Knox (1870). 


Phyllophichthus Gosline 


Phyllophichthus Gosline 1951a: 316. (Type spe- 
cies; P. xenodontus Gosline 1951a, by original 
designation.) 


DESCRIPTION: Body moderately elongate, cylin- 
drical, laterally compressed posteriorly; body = 
tail; snout produced, subconical, grooved on 
underside; lower jaw included; eye large; an- 
terior nostril subtubular, its posterior borders ex- 
tending downward into leaflike appendages, pos- 
terior nostril opens into mouth; GO vertical, 
crescentic, shorter than isthmus; dorsal and anal 
fins low, expanded before the sharply pointed 
tail tip; DFO above GO; pectoral fin well de- 
veloped; pop® absent; LL canal ossicles are short 
coils separated at pores; teeth conical, minute, 
and uniserial in jaws, recurved on mandible, ab- 
sent from vomer; neurocranium elongate, sub- 
truncate posteriorly, orbit large; PO strut absent; 
nasals and nasal cartilage well developed; SOC 
absent; maxilla fragile and not produced, articu- 
lated ca. mid-vomer (fig. 16); preopercle re- 
duced; suspensorium anteriorly inclined, jaw 
angle ca. 100°; otic bulla moderately developed; 
PG slender, short, and pointed anteriorly; hyoid 
slender, HH absent, CH with a minute anterior 
condyle (possibly a remnant of a fused HH2); 
branchiostegal rays numerous, filamentous, free 
from hyoid; UH split anteriorly, produced pos- 
teriorly as a slender spike; gill arches weak, 
pharyngeal teeth minute, C; absent, Hi ossified 
only at tip; pectoral girdle well developed, acti- 
nosts present; precaudal vertebrae more num- 
erous than caudal; coloration uniform. 


ETYMOLOGY: From the Greek P vAroY (phyl- 
lon), leaf, and Ophichthus (masculine), a genus 
of ophichthids, in reference to the leaf-like an- 
terior nostrils. 


DISTRIBUTION: Two described Pacific species, 
P. xenodontus Gosline from the central and west- 
ern Pacific and Indian oceans, and P. macrurus 
McKay*, described from Australia. 


Pisodonophis Kaup 
Pisodonophis Kaup 1856a: 47. Also spelt Piso- 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


dontophis by other authors. (Type species; 
Ophisurus cancrivorus Richardson 1844, as 
restricted by Bleeker 1865.) 

Pisoodonophis Kaup 1856b: 17. Emend. pro 
Pisodonophis Kaup 1856a. 

Brachycheirophis Fowler 1944: 190. (Type spe- 
cies; Pisodonophis daspilotus Gilbert, in Jor- 
dan and Evermann 1898, by original designa- 
tion.) 


DESCRIPTION: General characteristics those of 
Ophichthus. Differences include: pectoral fin 
broad-based; pop® present or absent; teeth mol- 
ariform or granular, multiserial on jaws and vo- 
mer, largest at intermaxillary with a short gap 
separating those of the vomerine block; PO strut 
absent; maxilla elongate, slender posteriorly; 
innermost branchiostegal rays free from hyoid; 
teeth of pharyngeal plates generally small, pave- 
ment-like; pectoral girdle reduced, actinosts ab- 
sent, Sc absent in one species; coloration gen- 
erally uniform, spotted in one species. 


ETYMOLOGY: From the Greek 77 © oof 
(pisos), pea, °o gwovys (odons), tooth, and 
Oo Pry (ophis; masculine), serpent. 


DISTRIBUTION: Known from 8-10 species, a 
circumtropical genus. Included are: Ophisurus 
cancrivorus Richardson (IP), O. boro Hamilton- 
Buchanan (IP), O. hijala Hamilton-Buchanan 
(IP)*, O. hoevenii Bleeker (IP), O. hypselopterus 
Bleeker (IP)*, O. semicinctus Richardson (EA)*, 
Pisodonophis copelandi Herre (IP), P. daspilotus 
Gilbert (EP), Pisoodonophis zophistus Jordan and 
Snyder (IP) (=P. cancrivorus?), Ophichthys dro- 
micus Gunther (EA)*. 


REMARKS: The relationships of Pisodonophis, 
Myrichthys, and their Ophichthus-like ancestors 
are discussed in the remarks on Myrichthys. 

Omochelys Fowler, considered by recent au- 
thors to be a junior synonym of Pisodonophis, 
is here referred to Ophichthus. Omochelys cru- 
entifer is unlike the species of Pisodonophis in 
having few branchiostegal rays, all of which are 
attached to the hyoid. 

Ganguly and Nag (1964) described the func- 
tional morphology of the pectoral girdle and the 
acranial myomeric musculature of a fish they 
identified as Ophichthus boro. The pectoral 
girdle they have crudely illustrated is clearly not 
that of Pisodonophis boro, and is probably from 
a species of Ophichthus. Tilak and Kanji (1967) 
noted their error and properly described the 
morphology of the pectoral girdle of Pisodono- 


83 


phis boro in relation to its habit. 
Pogonophis Myers and Wade 


Pogonophis Myers and Wade 1941: 78. (Type 
species; P. fossatus Myers and Wade 1941, by 
original designation.) 


DESCRIPTION: Body moderately elongate, cylin- 
drical, laterally compressed posteriorly; body 
slightly longer than tail; snout blunt; jaws sub- 
equal, a single pendulous barbel on upper lip; 
eye large; anterior nostril tubular, with a pos- 
terior tag; DFO behind GO; pectoral fin minute; 
tail tip pointed; preopercular pores vertically, 
not obliquely aligned, pop*® absent; LL ossicles 
nearly continuous; teeth pointed, uniserial ex- 
cept on maxillae where biserial, largest at inter- 
maxillary which is continuous with those of vo- 
mer; skull subtruncate posteriorly, orbit large; 
PO strut absent; nasals and nasal cartilage re- 
duced; SOC weakly pointed; maxilla elongate, 
slender posteriorly; suspensorium posteriorly in- 
clined, jaw angle ca. 80°; otic bulla moderately 
developed; PG broad posteriorly, free and terete 
anteriorly; hyoid slender; HH separated from 
CH by a broad gap; branchiostegal rays numer- 
ous, all slender and associated with hyoid, the 
anteriormost 3-4 rays fused basally; UH broad 
anteriorly, short and slender posteriorly; _ gill 
arches reduced, C; absent, UP3-UPs separate; 
pectoral girdle weakly developed, actinosts ab- 
sent; caudal = precaudal vertebrae; coloration 
strongly spotted. 


ETYMOLOGY: From the Greek Tf WyYwWwyv 
(pogon), beard, and 6 ees (ophis; 
masculine), serpent, in reference to the consipcu- 
ous barbel on the snout. 


DISTRIBUTION: A single eastern Pacific species, 
ranging from the Gulf of California to Peru (as 
Ophichthus afuerae Hildebrand). 


Quassiremus Jordan and Davis 


Quassiremus Jordan and Davis 1891: 622. (Type 
species; Ophichthus evionthas Jordan and 
Bollman 1889, by original designation.) 


DESCRIPTION: Body moderately elongate, cylin- 
drical, laterally compressed posteriorly; body 
slightly longer than tail; snout developed, coni- 
cal, a short groove on underside; lower jaw in- 
cluded; eye large; DFO behind GO; pectoral fin 
minute; tail tip pointed; pop*® present; LL os- 
sicles continuous; teeth pointed, uniserial, larg- 
est at intermaxillary and continuous with those 


84 


of vomer; skull truncate posteriorly, orbit large; 
PO strut absent; nasals well developed, nasal 
cartilage weakly developed; SOC strongly point- 
ed; maxilla elongate, slender posteriorly; oper- 
cular margins entire; suspensorium anteriorly in- 
clined, jaw angle ca. 100°; otic bulla well de- 
veloped; PG broad posteriorly, free and terete 
anteriorly; hyoid slender, HH separated from CH 
by a broad gap: branchiostegal rays numerous, 
all slender and associated with hyoid, the an- 
teriormost 3-4 rays fused basally; UH broad an- 
teriorly, short and slender posteriorly; gill arches 
reduced, Cs absent, UP3;-UPs separate; pectoral 
girdle reduced to a slender Cl and SCI; caudal 
= precaudal vertebrae; coloration strongly spot- 
ted. 


ETYMOLOGY: From the Latin quassus, obliter- 
ated, and ramus (masculine), oar, in reference 
to the minute pectoral fins. 


DISTRIBUTION: Known from 3 New World spe- 
cies. Included are: Ophichthus evionthas Jordan 
and Bollman (EP), Ophichthys nothochir Gilbert 
(EP) (including Q. parvipinna Seale), and Quas- 
siremus productus Seale (WA) (including Q. gos- 
lingi Beebe and Tee-Van). 


REMARKS: The osteological description of Quas- 
siremus is based on Q. nothochir. Q. evionthas, 
the generic type, was examined from radiographs 
and gill arch dissection. 


Scytalichthys Jordan and Davis 


Scytalichthys Jordan and Davis 1891: 635. De- 
scribed as a subgenus of Mystriophis Kaup. 
(Type species; Ophichthys miurus Jordan and 
Gilbert 1882, by original designation.) 


DESCRIPTION: Body cylindrical, moderately 
elongate, longer than tail; snout very short, flat, 
subconical; jaws elongate, lower jaw inferior; 
eye moderate; anterior nostril in a short tube, 
posterior opens into mouth; GO _ low lateral, 
crescentic, longer than isthmus; DFO well be- 
hind pectoral tips; pectoral fin reduced; tail tip 
bare, sharply pointed; pop® absent; LL ossicles 
continuous, not separated at pores; teeth strong, 
conical, those on maxilla in two widely separated 
rows, uniserial on dentary; those on vomer large, 
uniserial, widely separated, and continuous with 
those on intermaxillary; skull subtruncate pos- 
teriorly, orbit reduced; PO not bracing maxilla; 
nasals very reduced, nasal cartilage reduced; 
SOC absent, a short broad posterior projection; 
maxilla elongate, bifurcate posteriorly, the distal 


SERIES 4, V. 41, #1 McCOSKER — EELS 


section elongate and terete, the medial shorter 
and pointed; opercular series well developed, 
their margins entire; suspensorium nearly verti- 
cal, hyomandibular strongly ridged; otic bulla well 
developed; PG slender, a fine point anteriorly; 
hyoid arch slender, HH widely separated from 
CH; branchiostegal rays numerous, all slender, 
unbranched, and along arch; UH a short spike 
posteriorly; Cs short, UP3-UPs nearly fused, lower 
pharyngeal plate elongate; pectoral girdle re- 
duced, SCI and actinosts absent; IM bones, ribs, 
and CTP well developed; caudal vertebrae fewer 
than precaudal; coloration strongly spotted. 


ETYMOLOGY: From the Greek GZKUTAKANR 
(skytale), viper, and cyous (ichthys; mas- 
culine), fish. 


DISTRIBUTION: A single eastern Pacific species, 
ranging from the Galapagos Islands to the Gulf 
of California and Guadalupe Island. 


REMARKS: Schultz (1942) has commented on 
the generic status of Scytalichthys with regard to 
it dentition. 


Xyrias Jordan and Snyder 


Xyrias Jordan and Snyder 1901: 864. (Type spe- 
cies; X. revulsus Jordan and Snyder 1901, by 
original designation.) 


DESCRIPTION: Body cylindrical, moderately 
elongate, longer than tail; snout short, subconi- 
cal; jaws elongate, the lower inferior; eye mod- 
erate; anterior nostril non-tubular, laterally ovate, 
with a minute ventral flap; posterior nostril out- 
side mouth and covered with a flap; GO low lat- 
eral, crescentic, longer than isthmus; DFO be- 
hind pectoral tips; pectoral fin moderate; tail 
tip bare, pointed; pop* absent; teeth strong, 
conical, not extremely enlarged; those of vomer 
largest, uniserial, and nearly continuous with 
those of intermaxillary, jaw teeth multiserial; 
orbit depressed; PO not bracing maxilla; nasals 
and nasal cartilage reduced; SOC with a short 
posterior projection; maxilla elongate, bifurcate 
posteriorly; opercular series well developed, 
their margins entire; suspensorium nearly verti- 
cal; PG slender, tapering to a fine point an- 
teriorly; hyoid arch moderately slender, HH sep- 
arated from CH by a gap; branchiostegals num- 
erous, slender, unbranched and along arch; C; 
ossified, UPs-UPs united by a suture; pectoral 
girdle complete, SCI, Cl, Sc, Co, and actinosts 
visible in radiograph; IM bones, ribs, and CTP 
well developed; caudal vertebrae fewer than pre- 
caudal; coloration of numerous small spots. 


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ETYMOLOGY: From the Greek JuPioy 
(masculine), a shaveling, in reference to the ab- 
sence of cirri on the lips. 


DISTRIBUTION: A single species from Japan. 
Glover (1973) reported that it is caught occasion- 
ally by shrimp trawlers in the Misaki, Kochi, and 
Kumano prefectures. 


REMARKS: This description was prepared from 
radiographs and a gill arch dissection of the 
holotype. 


Comparison with Previous Classifications 


The results of the present study offer certain 
insights into the reliability of superficial morpho- 
logical characters as a basis for classification. The 
most recent attempt at a compilation of genera 
within the Ophichthidae was that of Rosenblatt 
and McCosker (1970). That study was based on 
an analysis of previous literature and an exami- 
nation of all recognized genera except Neen- 
chelys, Malvoliophis, and Benthenchelys (Castle, 
1972, was the first to suggest that Benthenchelys 
is an ophichthid). The following discrepancies ex- 
ist between the results of that study and the 
present: 

(a) Echelus was included in the Myrophinae 
(as Echelinae), following Gosline (1952); in the 
present study Echelus is shown to be an ophich- 
thine, and the presence of a caudal fin is not 
regarded as a definitive character separating the 
ophichthid subfamilies; 

(b) Leptenchelys was included in the Myro- 
phinae, again based on the caudal condition; in 
the present study it is suggested that its rela- 
tionships lie with the Bascanichthyini; 

(c) Lamnostoma was considered to be syn- 
Oonymous with Caecula; in the present study 
osteological characters have been identified that 
allow its generic separation; 

(d) Omochelys was considered to be synony- 
mous with Pisodonophis on the basis of tooth 
characters; in the present study it is indicated 
that its affinities lie closer to species of Ophich- 
thus than of Pisodonophis, and Omochelys is 
tentatively placed in the synonymy of Ophich- 
thus awaiting a revision of that large and cum- 
bersome genus; 

(e) Jenkinsiella, and Microdonophis and Zono- 
phichthus, were recognized at the generic level; 
in the present study no osteological characters 
have been found to support their generic recog- 
nition, and they are reduced to subgeneric rank 
under Cirrhimuraena and Ophichthus, respec- 


85 


tively. 

The results of this comparison however, have 
shown that the previous classification, based for 
the most part on external morphological char- 
acters, was not seriously upset by a classification 
based primarily on osteology. The finding of 
greatest Consequence was that the affinities of 
Echelus lie with the Ophichthinae rather than 
the Myrophinae; the caudal fin condition was 
shown to be trivial when compared with the 
“Ophichthus-like’’ condition of numerous osteo- 
logical characters. 

In that the majority of classification schemes 
within the teleosts are now based on external 
morphological characters, it is somewhat reas- 
suring to realize that at least within the Ophich- 
thidae, the external morphological classification 
parallels that based on osteology. 


EVOLUTION OF THE OPHICHTHIDAE 
Relationship to other Anguilliforms 


Recent authors have summarily recognized 23 
families within the suborder Anguilloidei (Green- 
wood, et al., 1966; Gosline, 1971). Subsequent 
studies have changed this listing in the following 
manner. Robins and Robins (1970) have ex- 
panded the Dysommidae to include the Dysom- 
minidae and the Nettodaridae and (1971) pro- 
visionally united the Nessorhamphidae with the 
Derichthyidae. Smith (1971) has provisionally re- 
moved Coloconger from the Congridae and 
erected the family Colocongridae. The above- 
mentioned authors have allocated the anguilloid 
families to five superfamilies, namely: the An- 
guilloidea (those eel families with paired fron- 
tals), Synaphobranchoidea (Synaphobranchidae, 
Dysommidae and Simenchelyidae), Congroidea 
(Congridae, Colocongridae, Muraenesocidae, Net- 
tastomidae, and Macrocephenchelyidae), Nem- 
ichthyoidea, and Ophichthoidea. 

The superfamiliy Ophichthoidea is restricted 
to the Ophichthidae. They appear most closely 
related to the superfamily Congroidea, but differ 
in the condition of several major characters (sev- 
eral fundamental characteristics of the Ophich- 
thidae and related apodal families are indicated 
in Table 7). As Smith (1971) has shown, the con- 
groids possess a complete pterygoid arch and a 
nearly complete gill arch skeleton. Conversely, 
the ophichthid pterygoid is reduced and. well 
separated from the vomer, and their gill arches 
exhibit various stages of reduction. The ophich- 
thids also differ from the congrids in possessing 


86 


numerous and overlapping branchiostegal rays, 
a fused frontal commissure, and a cartilaginous 
connection between the first epibranchial and 
second infrapharyngobranchial of the gill arch 
skeleton. Certain congrids, primarily within the 
subfamily Heterocongrinae, are similar to oph- 
ichthids in the development of laminar ribs, re- 
duced neural spines, an elongate body, and a 
fleshy tail tip (cf. Bohlke, 1957; Rosenblatt, 1967). 
Rosenblatt discussed these similarities and sug- 
gested (p. 95) that ‘the superficial similarties be- 
tween Gorgasia and the ophichthids are cer- 
tainly parallel adaptations to a similar mode of 
life, and the minor osteological similarities may 
be adaptations as well.” 


The general condition of the primitive ophich- 
thid genera, viz., Echelus and Ophichthus, ap- 
pears to be derived from a congrid ancestor. To 
my knowledge, no known congrid exhibits defin- 
itive ophichthid characters in an intermediate 
state of development, particularly the fused 
frontal commissure and overlapping branchio- 
stegal rays. The achievement of the ophichthid 
condition may have been a quantum jump in 
apodal evolution, attained by either an advanced 
congrid or congrid ancestor by means of a rapid 
integration of the above mentioned characters. 
Yet this may only be presumed until further evi- 
dence from extant or fossilized species is ob- 
tained. 


SERIES 4, V. 41, #1 McCOSKER — EELS 


Evolution within the Ophichthidae 


The purposes of any classification system are 
to best reflect the phylogenetic history of the 
taxa involved and to provide predictability, such 
that a newly discovered taxon might be properly 
placed without upsetting the system. An evolu- 
tionary scheme may only be inferred from exist- 
ing data, and must carefully weigh the prob- 
abilities related to each hypothetical pathway. 
In dealing with supra-specific categories, one is 
faced with the difficulty of delineating groups 
on a subjective basis. The history of the Ophich- 
thidae, however, would suggest that the genus, 
tribe, and subfamily reflect certain biological 
realities in an evolutionary sense. The present 
study has attempted to define and identify these 
groupings. The presence of several fundamental 
ostelogical characters in both major lines of the 
Ophichthidae suggests a monophyletic origin of 
the family. These characters include the frontal 
commissure, the epibranchial interconnections, 
and the unique manner of branchiostegal over- 
lap. The two subfamilies differ trenchantly in the 
manner in which the branchiostegal rays are at- 
tached to the hyoid, yet this difference need not 
preclude a monophyletic origin. The inferred 
subfamilial and tribal evolution within the Oph- 
ichthidae is illustrated below in Figure 37. Gen- 
eric evolution within each tribe is treated in the 
following discussion. 


Callechelyini 


Sphagebranchini 


Ophichthini 


Bascanichthyini 


Bentnenchelyini 


/ ‘ 


, 
a Myrophini 


CONGRID-LIKE ANCESTOR 


Figure 37. Proposed evolutionary relationship of ophichthid tribes. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Sehtsmorhynehus (1) 
Sehultatdta (2) 
snout, nostril, and hyoid 
dentition and neurocranium modified 
reduced, pop? absent 


ee aes (ea 20) 


pectoral fin lost, girdle reduced, 
Ahlta (1) gill arches reduced, CTP developed 


vomerine teeth lost 
jaws modified 


Myrophts (9) Pseudomyrophis (2) 

pectoral fin and girdle 
reduced 

posterior epipleurals 
lost 


Neenchelys (2) 


posterior nostril 
labial 
_-Benthenchelys (1) 
anterior nostril non-tubular, 
neurocranium shortened 


anterior nostril tubular, posterior 
nostril before eye, pectoral fin well 
developed, scapula and coracoid present 


CONGRID-LIKE ANCESTOR 


Figure 38. Proposed evolution of the Myrophinae. Number of species in parentheses. 


87 


88 


Myrophini and Benthenchelyini. Two. condi- 
tions indicate that the Myrophinae probably 
arose from the first dichotomy of the ophichthid 
lineage (Figs. 37-38). These are the presence of 
a well developed caudal fin and the unique my- 
rophine branchiostegal condition in which the 
majority of the rays are free from and well be- 
hind the epihyal. The attached rays, like those 
of the congirds, are basally broadened. Although 
the species of Echelus possess a myrophine-like 
caudal fin, they also possess an ophichthine-like 
hyoid and other osteological conditions that link 
them with the Ophichthinae. 

Primitive conditions of certain characters with- 
in the Myrophinae include: 


(a) teeth present on the dentary, intermaxil- 
lary, maxilla, and vomer; 

(b) pectoral fin moderately to well developed; 

(c) coracoid and scapula present; 

(d) posterior nostril lateral; 

(e) first basibranchial ossified; 

(f) seven attached branchiostegals along epi- 
hyal; 

(g) anterior nostril tubular; 

(h) eye not enlarged, its diameter ten or more 
in head length. 


— 


Benthenchelys cartieri, the single species with- 
in the Benthenchelyini, is highly specialized for 
its unique adaptation to a pelagic existence (see 
Remarks on Benthenchelys). It has however, re- 
tained certain primitive conditions which would 
suggest that it separated early from the ancestral 
myrophine stock. Benthenchelys possess all the 
primitive conditions listed above except that its 
first basibranchial has been reduced to cartilage. 
Whether Benthenchelys separated before, after, 
or along with the Neenchelys-Pseudomyrophis 
lineage was not discerned from the available 
data. The posterior nostril of Neenchelys and 
Pseudomyrophis, like that of Benthenchelys, is 
lateral. The nostril of Benthenchelys is extremely 
atypical within the Ophichthidae in that it is 
markedly more dorsally located than it is in 
other genera. The partial re-ossification of the 
third hypobranchial in Benthenchelys probably 
represents a secondary specialization; the third 
hypobranchial of all congrids, and presumably 
that of the basal ophichthids, is cartilaginous. 


Similarities between Neenchelys and Pseudo- 
myrophis were described earlier in this study. 
These genera have retained the first basibranch- 
ial, the pectoral fin, and pectoral girdle, but an 
attached branchiostegal has been lost in two of 


SERIES 4, V. 41, #1 McCOSKER — EELS 


the three species examined. Pseudomyrophis has 
become specialized in the reduction of its pec- 
toral fin and girdle, and in one species the body 
has become extremely elongate. Neenchelys is 
somewhat more primitive in that it has retained 
a developed pectoral fin and its girdle is not 
reduced. 

Beyond the level of the Pseudomyrophis-Neen- 
chelys separation, the posterior nostril has be- 
come labial in position. In general, the ophich- 
thid posterior nostril lies within the mouth or 
along the lip and is covered by a flap. The rever- 
sion to the more generalized and presumably 
primitive condition in which the posterior nostril 
lies along the outer edge of the lip has been 
achieved numerous times. This is evidenced in 
the contrasting posterior nostril conditions of 
closely related genera such as I/Ichthyapus and 
Apterichtus, Ophichthus and Ophisurus, and par- 
ticularly between the species of Muraenichthys. 
A distinction however, should be made between 
the condition of the earlier-removed myrophine 
genera (Benthenchelys, Pseudomyrophis, and 
Neenchelys) and that of the remaining ophich- 
thids. In no case does it appear that the posterior 
nostril has secondarily returned to a congrid-like 
placement. 

The next major dichotomy is that of the Ahlia- 
Myrophis lineage. That they arose from a com- 
mon ancestor is evidenced by their conspicuous 
specialization in which the pleural ribs are lim- 
ited to the anterior trunk vertebrae. The species 
of Myrophis differ considerably in external ap- 
pearance, primarily in body and snout elonga- 
tion, even though they differ little osteologically. 
Ahlia egmontis, by comparison, is specialized in 
having lost the vomerine dentition, posteriorly 
shortened maxillae, modified pterygoids, the 
hypohyals either lost or fused to the ceratohyal, 
and the dorsal fin origin withdrawn to the level 
of the anus. 

The remainder of the Myrophini comprise the 
species of Muraenichthys, Schultzidia, and 
Schismorhynchus, commonly called the ‘worm 
eels’. The reductions and specializations of the 
latter two genera must preclude them as ances- 
tors to Muraenichthys or to each other. Within 
Muraenichthys however, are species sufficiently 
generalized that either Schismorhynchus or 
Schultzidia might have been derived from them. 
As discussed in the remarks on Muraenichthys, 
subgeneric lineages, although including rather 
divergent extremes, are bridged by a broad spec- 
trum of morphological conditions. The more gen- 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


eralized species of the subgenus Scolecenchelys 
are likely ancestors to the subgenus Muraenich- 
thys and the genera Schultzidia and Schismor- 
hynchus. Species of Scolecenchelys have the pos- 
terior nostril within the mouth, separate upper 
pharyngeal tooth plates, an ungrooved snout, 
uniserial or biserial conical teeth, and a third pre- 
opercular pore (pop*). Species of the subgenus 
Muraenichthys have an external posterior nostril, 
multiserial granular dentition, a deep body, a 
blunt snout, and fused upper pharyngeal teeth. 
The species of Schultzidia have lost the pop® and 
the dentition is extremely reduced, being absent 
on the vomer, absent or embedded on the inter- 
maxillary, and minute or villiform in the jaws. 
Schismorhynchus labialis is specialized in quite 
a different manner, and seems to have been de- 
rived from a Muraenichthys-lineage different 
from that of Schultzidia. It is an elongate worm 
eel that has undergone specializations apparently 
related to its mode of feeding. A prominent 
toothed groove divides the underside of the snout 
and extends anteriorly to the elongate tubular 
nostrils. The suspensorium is forwardly inclined, 
and although the dentition is generalized, the 
upper pharyngeal tooth plates are fused, and 
certain gill arch members (Hs and lz) are absent. 


Ophichthini. The Ophichthini include the most 
primitive and generalized of ophichthids. Primi- 
tive conditions of certain characters include: 


(a) caudal rays developed; 

(b) pectoral, dorsal, and anal fins well devel- 
oped; 

(c) branchiostegal rays fewer than 20; 

(d) teeth conical, often multiserial, present on 
maxilla, dentary, vomer, and intermaxillary; 

(e) fifth ceratobranchial (Cs) ossified as a 
slender rod, upper pharyngeal tooth plates sepa- 
rate; 

(f) third preopercular pore (pop*) present; 

(g) neurocranium truncate posteriorly; 

(h) tail approximately 60 percent of the total 
length. 


The species of Echelus possess all of the above 
characters and represent one branch of the oph- 
ichthine lineage (Fig. 39). All subsequent oph- 
ichthins are specialized in having the tail tip re- 
duced to a finless point. Although differences 
exist in the condition of the tail tip, in no in- 
stance are caudal rays developed as in Echelus 
or the myrophines. The genus Ophichthus, as 
broadly defined herein, is the most generalized 
of the remaining ophichthine genera, having 


89 


character states that embrace most of the varia- 
tion of the other genera. The changes in other 
genera have to do with the loss or specialization 
of characters, or rearrangements of the basic 
Ophichthus condition. Certain lineages within 
Ophichthus appear ancestral to lineages within 
the tribe itself. For example, those species with 
few branchiostegal rays and an average body 
taper probably gave rise to the long-jawed pi- 
scivorous genera. The proposed sphagebranchin 
lineage can also be derived from generalized 
ophichthin characters. 

Ophisurus appears to be an offshoot from a 
moderately elongate Ophichthus-like species 
with few branchiostegals, nearly uniform denti- 
tion, and well developed surface sensory papil- 
lae. Its jaws and neurocranium modifications 
merit its generic recognition. 

The species of Quassiremus share several prim- 
itive characters with Ophichthus, but have be- 
come specialized through the reduction of the 
pectoral fin and loss of the C; and pectoral girdle 
elements. 


The species of Pisodonophis, Myrichthys, and 
Cirrhimuraena probably arose from a common 
lineage, evidenced in their increased number of 
branchiostegals and generally multiserial denti- 
tion. Cirrhimuraena is specialized in the devel- 
opment of labial cirri. Differences between the 
species of Cirrhimuraena are probably deserving 
of subgeneric rank, as discussed in the remarks 
on this genus. Pisodonophis and Myrichthys 
share several unique simlarities, primarily their 
multiserial molariform dentition and broad-based 
pectoral fins. Myrichthys is further specialized in 
the loss of pop’, the reduction of the pectoral fin 
and girdle, and the advancement of the DFO. 
As mentioned in the remarks on Myrichthys, a 
nearly continuous character series exists from 
species of Ophichthus —> Pisodonophis > 
Myrichthys. 

The species of Mystriophis, Echiophis, Brachy- 
somophis, Aplatophis, Xyrias, and Scytalichthys 
form a natural group of predaceous ophichthines 
specialized for the capture of large struggling 
prey. These specializations include the develop- 
ment of a postorbital strut to brace the maxilla, 
the strengthening of the suspensorium, enlarge- 
ment of the vomerine and jaw dentition, the ad- 
vancement and dorsal location of the eyes on 
the snout, and the attainment of a large size as 
adults. Certain species of Ophichthus (O. ophis, 
O. triserialis, and O. zophochir) are similarly 
specialized. The character states of Echiophis 


McCOSKER — EELS 


SERIES 4, V. 41, #1 


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PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


and Mystriophis are merely further specializa- 
tions of the condition of those species of Oph- 
ichthus. As mentioned in the remarks on Mystri- 
ophis, the specializations of EF. blastorhinos, when 
further investigated, may merit generic recogni- 
tion. Remaining genera in the lineage are further 
specialized by having a proportionately longer 
trunk region, or conversely, a shorter tail, and 
have lost the pop*. The examined species of 
Brachysomophis have developed labial cirri and 
a massive postorbital strut. The translocation of 
the orbit to the snout tip and development of 
the surface sensory papillae system in Brachy- 
somophis and Scytalichthys correlate with the 
feeding behavior; these species, like those of 
Echiophis (personal observation, and Hiatt and 
Strasburg, 1960), lie within their burrows with 
only their snout tips exposed, awaiting the pas- 
sage of prey items. The advantages of the eye 
position and papillae development are obvious. 
A similar feeding behavior is practiced by other 
ophichthines. Species of Callechelys, for ex- 
ample, have been observed (personal observa- 
tion, and R. H. Rosenblatt, personal communica- 
tion) with the head protruding from the sand 
bottom, apparently awaiting the passage of prey 
items. Callechelys has not undergone the devel- 
opment of the strengthened suspensorium and 
elongated jaws and associated orbit translocation 
typical of the long-jawed ophichthines, and feeds 
primarily on relatively smaller and weaker prey. 
The development of labial cirri in Brachysomo- 
phis and other ophichthine genera probably 
serves as a screen to prevent fine sediment from 
entering the mouth. The absence of the post- 
orbital strut in the remaining genera is assumed 
to be a secondary reduction. Xyrias and Scytal- 
ichthys are quite similar in the shape of their 
maxillae and in the condition of their multiserial 
dentition. Scytalichthys has a proportionately 
longer trunk region and has a further modified 
snout condition. Aplatophis is conditionally 
placed at the apex of this lineage. Its numerous 
reductions and specializations make its place- 
ment within the lineage difficult. 

The remaining ophichthine genera have lost 
the pop*. Malvoliophis has diverged from the 
Ophichthus condition in having an advanced 
DFO. Evips has retained an Ophichthus-like DFO, 
but has a considerably reduced pectoral fin and 
girdle and a proportionately reduced tail length. 
The ancestral bascanichthyin probably arose from 
this lineage, and probably possessed an anterior 
DFO, an ossified C;, and a reduced pectoral fin 


91 


and girdle. Subsequent ophichthins have lost the 
rod-like ossified C; possessed by all other oph- 
ichthins. Pogonophis, although similar in external 
appearance to species of Ophichthus, is special- 
ized in its development of labial barbels and by 
its loss of the C; and pop’. Leiuranus and Elapso- 
pis are closely-related genera which have de- 
parted from Ophichthus in snout shape, UP3-UP4 
fusion, and reduced pectoral fins and girdles; 
Leiuranus is further specialized in the loss of 
vomerine dentition and certain pectoral ele- 
ments. Phyllophichthus is aligned with this line- 
age although its exact placement is undeter- 
mined. Its suspensorium, jaws, dentition, and 
neurocranium are extremely modified. 


Sphagebranchini. The Sphagebranchini com- 
prise a specialized ophichthin offshoot of highly 
modified species, assembled on the basis of the 
absence of the pectoral fin, pectoral girdle reduc- 
tions, and low or entirely ventral gill openings. 
Primtive conditions of certain characters include: 


(a) neurocranium depressed; 

(b) branchiostegal rays fewer than 20; 

(c) pop* and tp? pores present; 

(d) Cs; ossified; 

(e) dorsal and anal fins present; 

(f) anterior nostrils tubular, posterior nostrils 
within mouth; 

(g) body and tail nearly subequal; 

(h) gill openings low lateral. 


The interpretation of intergeneric relationships 
within this tribe is made difficult by the reduc- 
tion or loss of numerous characters. Several in- 
terpretations are possible, depending upon the 
importance applied to certain characters. The 
following interpretation, in assuming the tribe to 
have had a monophyletic origin, assumes that the 
C; has been lost independently in two lineages. 
This loss seems plausible through an ossified > 
cartilaginous transformation, and has apparently 
occurred elsewhere in the family. 

The preliminary dichotomy separates the three 
sharp-snouted, entirely finless genera (Fig. 40). 
Each possesses a peculiar projection from the 
parapophyses of the anterior trunk vertebrae 
(Fig. 33A), a specialization not observed else- 
where in the tribe. Cirricaecula is primitive in 
having retained an Ophichthus-like C; but spe- 
cialized in its UPs3-UPs fusion, entirely ventral 
non-converging gill openings, and labial cirri de- 
velopment. The species of /Ichthyapus have a 
small eye, reduced anterior nostril, and cartilagi- 
nous C;. Species of Apterichtus have entirely lost 


McCOSKER — EELS 


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PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Caralophia (1) 
‘\ 


antervor nostril and snout 
modified, dentition reduced 


Phaenomonas (2) 

Ethadophts (2) 

tins, reduced tO a short. dorsal, 
trunk extremely elongate 


Gordtichthys (1) Leptenchelys (1) 
Cart tip) fieshy 


pectoral fin absent 


Agttpe (1) 


DFO posteriorly 
displaced 


to CALLECHELYINI 


a 


Bascanichthys (ca. 16) 
Co lost 


C. reduced or absent, 
DFO advanced to head, 
trunk elongated 


Dalophis (5)=——=——? 


OPHICHTHIN-LIKE ANCESTOR 


Figure 41. Proposed evolution of the Bascanichthyini. 


93 


94 


the Cs; and the posterior nostril is translocated to 
the outer lip. 

The remaining genera are characterized by an 
increased number of branchiostegals and the loss 
of the C;. They may be separated into two major 
lineages. Caecula and Stictorhinus are similar to 
the finless sphagebranchins in having ventral gill 
openings, pointed snouts, and moderately de- 
pressed neurocrania; these similarities, in part, 
may be adaptations to a similar mode of life. 
The derivation of Yirrkala and the closely-related 
Achirophichthys and Lamnostoma is somewhat 
uncertain. They have lost the pop® and have low 
lateral to ventral gill openings. The neurocrania 
of these genera are considerably modified from 
the broad, depressed state of other sphagebran- 
chins. The neurocranium of Yirrkafa is rounded 
dorsally. That of Lamnostoma has become. nar- 
rowed along the ethmoid, in probable correla- 
tion with its enlarged dentition and related to its 
feeding mode. Certain species of Yirrkala have 
retained the tp®. The species of Yirrkala are gen- 
erally elongate whereas those of Achirophichthys 
and Lamnostoma are cylindrical and stout. 


Bascanichthyini. The Bascanichthyini, like the 
Sphagebranchini, appear to be derived from a 
moderately specialized ophichthin-like ancestor. 
Certain specializations in the form of hyoid mod- 
ifications and fin reductions had already been 
achieved by their supposed ophichthin-like an- 
cestor. As adults, the bascanichthyins have spe- 
cialized toward a burrowing, vermiform existence, 
feeding on small prey and rarely leaving the 
substrate. As noted in the discussion of the axial 
skeleton (p. 45), the trunk elongation of species 
of Phaenomonas, Allips, certain Bascanichthys 
and presumably Gordiichthys is a specialization 
for this mode of life. Primitive conditions of cer- 
tain characters within the Bascanichthyini in- 
clude: 


(a) pectoral fin rudimentary; 

(b) eye small, but not minute; 

(c) Cs present, but reduced; 

(d) body and tail nearly subequal; 

(e) branchiostegal rays numerous; 

(f) DFO behind head; 

(g) snout grooved on underside; 

(h) gill openings low lateral in position. 

The inclusion of Dalophis in the Bascanichthy- 
ini is uncertain. Although possessing Ophichthus- 
like body/tail proportions and an ossified C;, 
it appears more similar to the bascanichthyin 
condition in its fin reductions and general ce- 


SERIES 4, V. 41, #1 McCOSKER — EELS 


phalic appearance. As a_ bascanichthyin, it is 
placed near the primary separation from the 
ancestral lineage (Fig. 41). 


The remaining genera appear to be derived 
from the generalized conditon of Bascanichthys. 
The rudimentary pectoral fin is retained only by 
Allips. Allips is similar to species of Bascanich- 
thys in its trunk elongation and general physi- 
ognomy, but differs in its presumably secondar- 
ily-derived posterior DFO. Phaenomonas is fur- 
ther specialized through reduction from this line- 
age and has become nearly finless. Gordiichthys, 
not examined in this study, is provisionally re- 
ferred to this lineage on the basis of characters 
included in its terse description. The remaining 
bascanichthyins are somewhat similar in their 
general morphologies, and have undergone sev- 
eral modifications and reductions from the Bas- 
canichthys condition. The Callechelyini are here- 
in assumed to be derived from a bascanichthyin- 
like ancestor, but have further specialized and 
radiated along a different complex of adaptive 
characters. 


Callechelyini. The Callechelyini is the most 
distinct and compact of ophichthine tribes. The 
species are among the most specialized of oph- 
ichthids and are quite removed from the ances- 
tral ophichthid stock. Their specializations and 
reductions impart a particular facies to the group 
that readily separates it from other ophichthids, 
particularly evidenced in the ventral, convergent 
gill openings, laterally compressed body, short 
tail, anterior dorsal fin origin, small eyes ,and 
reduced pore systems. These outer similarities 
are borne out by the shortened neurocranium, 
stout hyoid, and the osteological reductions in 
the pectoral apparatus and gill arches which 
further characterize the group. 


These conditions appear to have been de- 
rived from a bascanichthyin-like ancestor or from 
an ophichthin ancestor which gave rise to the 
Bascanichthyini. Available specimens or radio- 
graphs of 20 of the 22 species of the Callechely- 
ini have allowed an in-depth study of this tribe. 
The meristic and morphological characters listed 
in Tables 8-9 were used to generate the com- 
puter-programmed taxonomic evaluations illus- 
trated in Figures 43-44. The characteristics of 
computer programs WVGM and REGROUP are 
described in the taxonomic methods sections of 
this study. 


Primitive conditions of certain key characters 
within the Callechelyini include: 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 95 


Letharehus (3) 


aaterior nostrils lost, 
/m supraorbital pore added, 
neurocranium elongated 


Paraletharechus (2) 
Calleehelys (15) 


UH forked, 
scapula present 

anak tin lost, branchial pouches UH spiked, 
expanded scapula absent 


—- =— 
nS 
‘ 


Aprognathodon (1) slender branchiostegals 
\ 


intermaxillary teeth lost, 


‘ 

\ 

gill arches modified ; 
snout grooved , 


/ 
/ 
/ 
Leuropharus (1) 


vomerine teeth lost 


EH branchvosteeals broad, rays 20-25), 
snout not grooved on underside, 
trunk not jexcessavely ellongate 


BASCANICHTHYIN-LIKE ANCESTOR 


Figure 42. Proposed evolution of the Callechelyini. 


96 


(a) teeth uniserial and present on the dentary, 
intermaxillary, maxilla, and vomer; 

(b) underside of snout not grooved; 

(c) gill openings low and bascanichthyin-like, 
not specialized as in Letharchus and Paralethar- 
chus; 

(d) branchiostegal rays fewer than 25 pairs, 
those along the epihyal broadened basally; 

(e) three supraorbital pores; 

(f) trunk not extremely elongate, tail 40 per- 
cent or more of total length; 

(g) urohyal ossified and spike-like, not forked 
posteriorly; 

(h) two longitudinal rod-like pectoral elements. 


No living tribal member possesses all the prim- 
itive characters listed above. However Aprogna- 
thodon platyventris and certain species of Cal- 
lechelys, except for minor specializations, closely 
approximate the above conditions. 

The retention of several primitive characters 
suggests that Aprognathodon platyventris sepa- 
rated early from the basal stock (Fig. 42). It has 
become specialized through the loss of inter- 
maxillary dentition and the incomplete re-ossi- 
fication of the third hypobranchial. These condi- 
tions are probably adaptations to a specialized 
mode of feeding. 

The next dichotomy in the tribal evolution in- 
volved two other New World genera, Letharchus 
and Paraletharchus (compare Figs. 42-44). They, 
like Aprognathodon, have retained the broad 
branchiostegals and ungrooved snouts, but are 
specialized in having broadly flared branchial 
pouches and in the loss of the anal fin. Species 
of Paraletharchus appear externally quite similar 
to species of Callechelys; species of Letharchus 
are quite distinctive in having lost the tubular 
anterior nostril condition, added a fourth supra- 
orbital pore, and having more elongate and de- 
pressed neurocrania. 

The remaining callechelyins include Leuro- 
pharus lasiops and the numerous species of 
Callechelys. Leuropharus is somewhat general- 
ized in having few vertebrae, a moderate tail 
length, and an ungrooved snout, yet it differs 
from other callechelyins in lacking teeth on the 
vomer. Whether it separated before the species 
of Callechelys or from a Callechelys-like ancestor 
is questionable in that L. Jasiops appears to have 
numerous, slender branchiostegals (observed 
from a radiograph of the type specimen). This 
condition, if observed correctly, is more ad- 
vanced than that of certain species of Callech- 
elys (C. nebulosus, C. springeri, and C. holo- 


SERIES 4, V. 41, #1 McCOSKER — EELS 


chromus), and would necessitate the parallel 
evolution of this condition if Leuropharus sepa- 
rated earlier than Callechelys. 

The remaining genus, Callechelys with 15 rec- 
ognized species, has apparently combined the 
generalized callechelyin condition with certain 
minor specializations into a very successful ‘“‘body 
plan’. Evolution within Callechelys, as discussed 
on page 63,has included two or three early 
separations from the ancestral stock, which were 
followed by two major subgeneric lines. Callech- 
elys nebulosus, C. springeri, and perhaps C. holo- 
chromus have retained broadened branchiostegal 
rays, which is probably correlated to their mod- 
erate number. These two conditions are probably 
responsible for the separation of these species 
from other Callechelys species in programs 
WVGM and REGROUP (Figs. 43-44). These con- 
ditions should not preclude their inclusion within 
Callechelys and _ illustrates the weakness of a 
numerical taxonomic scheme based on too few 
characters. The remaining species are special- 
ized in having more numerous and _ slender 
branchiostegal rays, and a longer trunk region 
and an increase in the number of trunk verte- 
brae. These species have evolved along two line- 
ages, one containing species which have retained 
a simple urohyal and lost the posterior pectoral 
girdle element (the scapula?), and another with 
species in which the urohyal is split posteriorly 
into two slender divergent rays but with the sec- 
ond pectoral element retained. Neither of these 
specializations, when compared with those of 
other ophichthids, seems to merit generic sepa- 
ration. 


Zoogeography and Comments on Ophichthid 
Speciation 


In the absence of a suitable fossil record, it is 
virtually impossible to reconstruct with certainty 
the past distribution and center of origin of the 
Ophichthidae. Certain inferences relating to an- 
cestral distributions however, can be made based 
on the present species distribution, assumed en- 
vironmental tolerances of ophichthids in general, 
and the presumed geological history of tropical 
land and water masses. 

Two major geological events have directly af- 
fected the distribution of tropical marine organ- 
isms. These were the Miocene (?) closure of the 
Tethyan Seaway through the convergence of the 
European and African continental plates (Phillips 
and Forsyth, 1972), isolating the Mediterranean 
and Atlantic from the Indo-Pacific, and the late 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 97 


Fab 


-1 .8 6 4 .2 0 .2 4 .6 .8 +1 
TEE RII eS 


A. platyventris 


L. velifer 
L.rosenbliatti 
P pacificus 
P.opercularis 
C.nebulosus 

L, lasiops 

C. bilinearis 

C. cliffi 
C.galapagensis 
Cc. muraena 

C. luteus 
Cspringeri 
Cmarmoratus 
Ceristigmus 
Cmelanotaenius 


Striatus 


Figure 43. Phenogram of the relationships of the species of the Callechelyini, using 
program WVGM. The levels of correlation at which species join are represented by the scale 


Tab- 


98 


Cis 


4/9 


C. springeri 


marmoratus 


Callechelys 
(9) 


SERIES 4, V. 41, #1 McCOSKER — EELS | 


8/9 


Letharchus 


(2) 


Aprognathodon 
2/3 
2/27 1/9 
Paraletharchus 
(2) ays Leuropharus 


nebulosus 


& C. 


Figure 44. Interrelationships of species groups of the tribe Callechelyini, as defined by 
program REGROUP. Significance level set at 0.600. Fractions are the ratios of the number of 
observed between-group species connections to the maximum number of possible connec- 
tions. Number of species represented per genus are within parentheses. Not included are 
Callechelys bitaeniatus, C. holochromus, C. leucopterus, and Letharchus aliculatus. 


Pliocene to Pleistocene closure (Whitmore and 
Stewart, 1965) of the Middle American Seaway, 
separating the New World oceans (Rosenblatt, 
1963). Assuming that the environmental toler- 
ances of the Ophichthidae have always limited 
them to tropical, sub-tropical, or warm temperate 
waters, it may be stated that the Tethyan and 
Central American Seaway closures have delimi- 
ted, in large part, the waters available to the 
distribution of living genera. (A single exception 
may be the distribution of Ophisurus serpens in 
the Mediterranean, and eastern and western Afri- 
can shores, probably resulting from a transgres- 
sion of the Cape of Good Hope.) Applying these 
assumptions to the known distribution of oph- 
ichthid genera (Table 10), inferences concerning 
the evolution and generic interrelationships of 
the family may be drawn. For example, the seven 
circumtropical genera must have existed prior to 
the closure of the Tethyan Seaway, or have 
passed through the Central American Seaway 


and transgressed both oceans. Recent informa- 
tion gained from investigations of plate tectonics 
and paleomagnetism suggests that the Atlantic 
Ocean during the Early Cretaceous was much 
narrower than at present (Phillips and Forsyth, 
1972). On that basis, the distribution of an ar- 
chaic species across the Tethyan Seaway and into 
the eastern Pacific would seem quite plausble. 

An Ophichthus-like genus probably existed in 
the Upper Eocene, as evidenced by Storms’ 
(1896) description of Eomyrus dolloi from the 
Wemmelian Formation of western Europe. The 
neurocranium, as illustrated, is much like that of 
a modern Ophichthus. Those genera restricted 
to both coasts of the New World (Table 10, 
Group III) must have existed prior to the Pana- 
manian uplift, and now include several pairs of 
closely related species. Echiophis is also a mem- 
ber of this group but has presumably extended 
its distribution to the eastern Atlantic. The genus 
Muraenichthys is presently limited to the Red 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Sea, Indian Ocean, western and central Pacific 
oceans, with a single south eastern Pacific spe- 
cies known only from the offshore Chilean is- 
lands of Juan Fernandez and San Felix. Its ab- 
sence from the tropical Atlantic, as explained by 
McCosker (1970), perhaps suggests a post-Teth- 
yan origin of the genus rather than the lack in 
the Atlantic of a suitable habitat. The single New 
World species is probably recently derived from 
the south-western Pacific by means of eastward 
transport across the South Pacific. The restriction 
of many genera to the Indo-Pacific and New 
World oceans (Table 10, Groups II, IV-V) prob- 
ably represents radiations since the Tethyan and 
Central American Seaway closures, respectively. 
The origin of genera with species distributed 
across major water masses might be _ inferred 
from an analysis of the species involved. The 
genus Phaenomonas, for example, contains an 
eastern Pacific species (P. pinnata) which ranges 
from the Gulf of California to Colombia, and an 
Indo-Pacific species (P. cooperae) which ranges 
from Hawaii to east Africa. The extreme speciali- 
zation of P. cooperae, in comparison to P. pin- 
nata, might suggest that Phaenomonas arose in 
the eastern Pacific and radiated westward. Its 
presumed absence from the Atlantic Ocean 
would indicate that Phaenomonas either evolved 
from the bascanichthyin stock since the closure 
of the Central American Seaway, or that proper 
Phaenomonas habitat is presently unavailable in 
the Atlantic. Another explanation might assume 
that the present existence of the primitive P. 
pinnata in the eastern Pacific is the result of 
an eastward radiation from an Indo-Pacific ori- 
gin, followed by further specialization of the an- 
cestral stock through competition with the more 
complex ophichthid fauna of the Indo-Pacific. 
Caution however, must be applied in any of the 
above assumptions concerning the present dis- 
tribution of ophichthid genera. Modern collect- 
ing methods have resulted in the discovery of 
numerous new species, many of which repre- 
sented new genera, and it is highly likely that 
many extant ophichthid species remain uncol- 
lected. 

An analysis of the distribution of species 
among ophichthid genera reveals several inter- 
esting biological phenomena. The classification 
proposed here is hopefully more than just a con- 
venient cataloguing system, and assumes that 
the disproportionate number of species in vari- 
ous ophichthid genera, ranging from a single 
species in nearly half of the genera to more than 


99 


50 in Ophichthus, reflects something real in the 
natural system. Figure 45 illustrates the distribu- 
tion of species among the genera of the Oph- 
ichthidae, Gobiesocidae, and the blenniid tribe 
Salariini. These examples were chosen because 
they represent recent monographic revisions, 
uncomplicated by the efforts of a multiplicity 
of authors; the gobiesocid data are from Briggs 
(1955) and subsequent species descriptions and 
the blenny data are based on Smith-Vaniz and 
Springer (1971). A pattern exists in the three 
groups illustrated, and if transformed to logs, the 
data would indicate an almost straight-line in- 
verse relation between the log-number of spe- 
cies/genus and the log number of genera. C. B. 
Williams (1964) has discussed similar evidence 
from a number of terrestrial groups and sug- 
gested that such a log-normal mathematical pat- 
tern is a recurrent phenomenon in natural sys- 
tems. He observed that the fit of these data to a 
calculated log series is moderately good at most 
levels, but higher than expected for monotypic 
genera. 


The following hypotheses are proposed to ex- 
plain the distribution of species among genera. 
One hypothesis might assume that such a dis- 
tribution of species among genera reflects the 
evolution of a group with many recently-derived 
genera, and a decreasing number of genera 
which have existed for increasingly longer time 
spans. Those archaic genera have had the op- 
portunity through geologic time and events to 
segregate and speciate, whereas the more recent 
taxa have lacked those opportunities. A more 
intriguing hypothesis however, suggested by 
Richard H. Rosenblatt (personal communication), 
might assume that a combination of characters 
exist in the ancestral lineage from which taxa 
radiate by means of specializations and reduc- 
tions; certain resultant taxa would include a 
combination of characters which would allow 
further radiation (dependent upon certain bio- 
logical factors and geological events) resulting in 
genera with numerous species, whereas other 
taxa have specialized in a manner which, in re- 
lation to the available environments, has a low 
probability of further radiation. The latter cate- 
gory contains the numerous monotypic genera 
of the Ophichthidae. These monotypic genera 
might represent evolutionary “forays’’ into rather 
unique environments or life styles and are ap- 
parently unsuccessful beyond their present lim- 
ited area of distribution or as ancestral bases for 
further speciation. The former category, in which 


SERIES 4, V. 41, #1 McCOSKER — EELS 


100 
25 
re) 
O0=Gobiesocidae 
20 A: Salariini 
% 
= O=: Ophichthidae 
¢ Oo 
0 15 
6 
= 
0 
5 10 fe) 
fe] A Oo 
P| 
A 
25 
O oO 
Oo Oo Opa 
A M A AAMOO DADA fe) 
bed 1 2 3 4 5 678910 20 30 40 5060 


Species /Genus Clog scale) 


Figure 45. Distribution of species among the genera of the Ophichthidae, Gobieso- 


cidae and Salariini. 


few genera contain many species, contains such 
genera as Myrichthys, Callechelys, and Oph- 
ichthus, in which the combination of adaptive 
characters selected for have, with minor modi- 
fications, resulted in the numerous species which 
occupy similar habitats in all tropical oceans. 
Biological factors such as the leptocephalus lar- 
val stage and geological events such as seaway 
closures lend credence to the first mentioned 


hypothesis and complicate the second. Yet the 
present distribution and few species of several 
presumably archaic ophichthid genera would 
tend to support the latter hypothesis. 

Further investigations into the ecology and 
behavior of species of this intriguing and diverse 
eel family may offer further insight into the evo- 
lutionary processes which have shaped _ the 
Ophichthidae. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


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75: 13-98. 


Trewavas, E. 1932. A contribution to the classifi- 
cation of the fishes of the order Apodes, based 
on the osteology of some rare eels. Proc. Zool. 
Soc. London, part 3: 639-659. 


Turton, W. 1807. The British fauna, containing a 
compendium of the zoology of the British 
Islands; arranged according to the Linnaean 
system. Swansea, London. 230 pp. (Not seen). 

Vahl, M. 1794. Beskrivelse af en nye fiskeslaegt. 
Skrivt. Naturh. Selsk. Kiobenhavn 3(2): 149- 
156. (Not seen). 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Wade, C. B. 1946. Two new genera and five new 
species of apodal fishes from the eastern Pa- 
cific. Allan Hancock Pac. Exped., 9(7): 181- 
DAS: 

Weber, M. and L. F. de Beaufort. 1916. The fishes 
of the Indo-Australian Archipelago. III. Ostario- 
physi: Il Cyprinoidea, Apodes, Synbranchi. 
Leiden. 455 pp. 

Whitley, G. P. 1930. Five new generic names for 
Australian fishes. Austr. Zool., 6(3): 250-251. 

Whitley, G. P. 1934. Studies in Ichthyology. No. 
8. Rec. Australian Mus., 19(2): 153-163. 

Whitley, G. P. 1940. Illustrations of some Aus- 
tralian fishes. Austr. Zool., 9(4): 397-428. 

Whitley, G. P. 1950. New fish names. Proc. R. 
Zool. Soc. New South Wales, 1948-1949: 44. 

Whitley, G. P. 1951. Studies in ichthyology. No. 
15. Rec. Australian Mus., 22(4): 389-408. 

Whitley, G. P. 1968. A check-list of the fishes 


107 


recorded from the New Zealand region. Austr. 
Zool., 15(1): 1-102. 

Whitmore, F. J., Jr. and R. H. Stewart. 1965. Mio- 
cene mammals and Central American seaways. 
Science, 148(3667): 180-185. 

Williams, C. B. 1964. Patterns in the balance of 
nature and related problems in quantitative 
ecology. Academic Press, London. 324 pp. 

Wormuth, J. H. 1971. The biogeography, system- 
atics and interspecific relationships of the 
oegopsid squid family Ommastrephidae in the 
Pacific Ocean. PhD diss., Univ. Calif., San 


Diego. 189 pp. 
Ypiranga Pinto, S. 1970. Observacoes ictiologicas. 
VI — Antobrantia, novo genero de ofictideo 


do Brasil (Actinopterygii, Anguilliformes, Oph- 
ichthyidae). Atas da Sociedade de Biologia do 
Rio de Janeiro, 14 (1-2): 13-15. 


108 


Table |. 


Abbreviations are: 
M, molariform; V, villiform. 
serial; U, uniserial. 


+, presents = 


Shape: 
Rows in jaws and vomer: 
I-V signifies intermaxillary-vomerine. 


SERIES 4, V. 41, #1 


Dentition of the Genera of Ophichthidae 


McCOSKER — EELS 


C, conical; F, fang-like; 


B, biserial; M, multi- 


Shape 


in Jaws Vomer 


I-V Gap 


Benthenchelyini 


Benthenchelys 


My rophini 
Ahita 
Muraent chthys 
Myrophts 
Neenchelys 
Pseudomyrophts 
Scehismorhynchus 
Schultztdia 


Callechelyini 
Aprognathodon| 
Callechelys 
Letharchus 
Leuropharus 
Paraletharchus 


Sphagebranchini 


Achtrophtchthys 
Apterichtus 
Caecula 
Cirriecaecula 
Hemerorhinus 
Ichthyapus 
Lamos toma 
Sttetorhinus 
Ytrrkala 


Bascanichthyini 


Alltps 
Bascanichthys 
Caralophia 
Dalophts 
Ethadophts 
Gorditchthys 
Leptenchelys 
Phaenomonas 


Ophichthini 
Aplatophtis 
Brachysomophis 
Cirrhimuraena 
Elapsoptis 
Echelus 
Echtophis 
Evips 
Leturanus 
Malvoltophis 
Myrichthys 
Mystriophis 
Ophtchthus 
Ophtsurus 
Phyllophtchthus 
Pisodonophis 
Pogonophts 
Quasst remus 
Seytaltchthys 
Xyrtas 


C,M 


(my Caz) (oe) (se) (se) 


(se) (sz) (sed se) Con) (ae) Ged ifs) aQan-e- 


eal 


GaGa 


(‘= 
Seo Sees Sse Sees 
o 


Sees eseSee 


SSeS iS SeSsesce 


See “eee 


crc 


U 


ie) 


Seste U Stet =giSg 


U,B 


te teterve¢t¢tr + 


tet te et 


1. Intermaxillary teeth absent 
2. Maxillary teeth biserial, dentary uniserial 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Table 2. Number and Location of Branchiostegal Rays of the Species of the 
Ophichthidae 


Counts represent the right side only. Rays joined basally are counted sepa- 
rately. ''CH-EH'' represents the cartilaginous CH-EH interspace. ''Free'' rays 
are noticeably separated from the hyoid. 


Total Free Along CH CH-EH Along EH 
Benthenchelys cartiert 20 13 = = 7 
Ahlia egmontts 47 4] - - 6 
Muraentchthys chilenstis 32 25 = = 7 
Muraentchthys gymopterus 43 36 = = 7 
Muraentchthys macropterus 31 37 = = 4 
Myrophis plumbeus 43 37 = = 6 
Myrophis uropterus 32 26 = = 6 
Myrophts vafer hg 42 - - 7 
Neenchelys buitendijki' 30 24 - - 6 
Pseudomyrophis micropinna 47 ho - - 7 
Pseudomyrophts ntmtus 23 17 = - 6 
Schismorhynchus labtalis 30 25 = = 5 
Schultzidia johnstonensts 33 29 - - 4 
Aprognathodon platyventris 28 = 14 2) 12 
Callechelys btlinearis 27] = 21 ] 5 
Callechelys ertstigmus 31 = 5 6 20 
Callechelys galapagensts 27 = 15 2 10 
Callechelys marmoratus 29 = 18 2 9 
Callechelys melanotaentus 29 = 12 3 14 
Callechelys nebulosus 29 10 8 3 8 
Letharchus rosenblattt 34 = 16 3 15 
Paraletharehus pact ficus 32 = 23 4 5 
Aptertchtus flavicaudus 16 l I l 13 
Caeeula pterygera 25 12 = 3 10 
Ctrrteaecula johnsoni 18 - ] 3 14 
Iehthyapus selachops 19 - 4 2 133 
Lamostoma ortentalis 29 26 = = 3 
Sttetorhinus potamius 29 - 5 4 20 
Yirrkala Lwnbricotdes 27 = 5 2 20 
Yirrkala misolensis 29 - 12 2 15 
Yirrkala tenuis 25 > 4 3 18 
Bascanichthys panamensts 30 ] 5 3 21 
Caralophia loxochila 27 = 8 4 15 
Phaenomonas cooperae 28 19 3 5 1 
Phaenomonas pinnata 26 19 1 6 = 
Aplatophis chaultodus 22 = 2 | 19 
Brachysomophts sauropsts 17 = 2 1 14 
Cirrhtmuraena macgregort 24 = 7 2 15 
Cirrhimuraena taentopterus 29 6 l 4 18 
Echelus myrus 15 = 4 e 1] 
Echelus pachyrhynchus 18 - 4 1 13 
Echtophis sp. 21 - 4 1 16 
Elapsopts cyclorhinus 24 4 2 1 17 
Leturanus semtctnctus 31 - 6 3 22 
Malvoltophts pinguts 21 = 4 1 16 
Myrichthys colubrinus 30 = 22 3 5 
Myrtchthys maculosus 35 - 9 2 24 
Myrtchthys xystrurus 31 3 3 ] 24 
Ophtehthus cephalazona 27 2 2 2 21 
Ophiehthus cruentifer 17 = 4 = 13 
Ophiechthus frontalis 19 = 5 ] 13 
Ophichthus triserialis 22 = 3 = 19 
Ophtchthus zophochir 27 = 5 1 21 
Ophisurus serpens 19 = 2 | 16 
Phyllophichthus xenodontus 29 22 = = 7 
Pisodonophts boro 31 I 6 2 22 
Pisodonophis canerivorus 30 6 | I 22 
Pisodonophis daspilotus 32 8 = 3 21 
Quasstremus nothochir 21 2 2 = 18 
Seytaltchthys miurus 25 - 4 2 19 


1. From Nelson (1966a: 


fig. 2a) 


109 


110 SERIES 4, V. 41, #1 McCOSKER — EELS 


Table 3. Gill Arch Condition in the Ophichthinae 


Abbreviations are: B,_y4, basibranchials; H1-3, hypobranchials; Cyi-c, ceratobran- 
chials; Io_ , infrapharyngobranchials; UP 3-4, upper pharyngeal dermal tooth plates; 
0, ossified; -O-, UP3-UP, fusion; C, cartilaginous; R, rudimentary; -, absent; *, 
from Nelson (1966a: Table 1) 


By Bo B3 Bh Hy Ho H3 C1-4 Cc 1) 13 UP3 UPL 
Ophichthus, zophochir o © &R © @ @ € 0 o © ®@ © 0 
0. rhytidodermatoides OC. Se 0 Oba 0 Yo © @ @ 0 
0. polyophthalmus* o ¢ Kk kK © @ € 0 OF O © 0) 
0. altiptnnis Ws eS Oo © f 0 Oo @ © © 0 
0. erabo yo ¢ & kK © © FE 0 OM oO © 0) 
O. eruentifer W es os = © @ ££ 0 oy O OM 0 
0. cephalazona 0 R > = ® @ 0 9 OG © @ 0 
Pisodonophis boro 2 Ss BR © oO —-E 0 XH @ 0) 0 
P. canertvorus 0 ny GG go @ © 0 OF 0) 0) 0 0 
Ophisurus serpens Oo © © & @ @ € 0) oO o OG © 0) 
Evips pereinetus 09 © FF kR O © CE 0 Oo © @ 0 
Echelus myrus oy ¢€¢ © CG O © 0 O00 0 0 
E. pachyrhynchus YO © sc © O © 0) J OO 0 0 
Myrichthys colubrinus Oo € = = ®% © CE 0 Oo O @ @ 0) 
M. maculosus 09 € = &€ O OM E 0 oO OM @ 0) 
M. xystrurus Onc = C 0) (0) C 0 00) 0h a0 0 
Aplatophis chaultodus QO «= =£ = M OO € 0) oO © @ 0 
Brachysomophts sauropsts oy CC = © @ O CE 0 oO @ @ 0 
B. henshawt* Y © oc & © 0 Oo © © @ 0 
Ctrrhimuraena macgregort Oo © R KR O O € 0 QO © ® © 0 
C. taentopterus ®O o> R > © © GE 0) 0 © @ 0 
Echtophis tnterttnetus* Oo f RF C OM OO CE 0) 09 © @ @ 0) 
Echtophis sp. 0 ¢ C Gc @ @ CC 0 oO =(0)= 
Xyrtas revulsus oO © = ¢€ @® @ EC 0 0) 0) 0 -0- 
Malvoltophis pinguts o © R KR O O CEC 0 Oo @ @ -0- 
Elapsopts cyclorhinus 9 € KR = O DO © 0 = @ -0- 
Leturanus semictnctus Oo €©€ Ss € GO © 0 = 0 =(0)= 
Phyllophtchthus xenodontus 0 CC R - O0O O C 0 = © @ @ 0 
Pogonophts fossatus 0 © = ss @ @ € 0 = @ @ 0) 
Quasstremus evtonthas 0 = - > (i Or (c 0 = 0) 0) (0) 0 
Q. nothochir QO =o = => © © G 0 = @ © 0 
Dalophis imberbts 0 Cc Kk © O @ 0) oo @) 0 
Bascanichthys teres* 9 © = € M O CE 0 QO © @ @ 0 
B. panamensis ) coc > € @O@ OM CE 0 >= @ @ © 0) 
Alltps concolor oo © = € OM © EC 0 = @ @ @ 0 
Ethadophts byrnet 0 € os = OM @ E 0 > ®}) @ 0 
E. merenda oO CC RR €C BO O CE 0 = © © 0 
Phaenomonas ptnnata Oo © = € O O CE 0 > 0 @ © 0 
Caralophia loxochila QO o © € OM © CEC 0 = 9 @ =(0)= 
Callechelys marmoratus 0) 6 >= @ @© € 0 > ®% @& @ 0 
Aprognathodon platyventris 0 C - - 0 0 0 0 >= 0 © @ 0 
Paraletharchus pact ficus 9 ¢€C = R © © EC 0 = 0 © © 0 
Letharchus velifer* Oo GF os f 6 @ 0 > @ © @ 0) 
Letharchus rosenblatti oy © = R @ © CEC 0) > @ © © 0 
Ichthyapus selachops Q@ oc = © ® © CE 0 QO @ @ @ 0 
Cirrieaeeula johnsont 06 C= = rR © © € 0 OO 0 =(0)— 
Apterichtus flavicaudus Q@ =e ces =s @® © € 0 = @ @ @ 0 
Caecula pterygera Oo © © € M ® FC 0 = @ © © 0) 
Yirrkala Lumbricotdes Ho CC C€ © 0 @ 0 = @ © @ 0 
Y. tenuts OF Re ac C One e0) 1G 0 >= @ © 0 
Y. misolensts oo € &€ & oO @ € 0 = @ © © 0 
Y. kaupt OC Ge Gc @ @ € 0 = 00) 0 0 
Lamostoma orientalis Oo © = = ® © Ef 0 2 9 @ @ 0 
Stictorhinus potamtus O € C¢ & © @ CF 0 = @ © © 0 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 111 


Table 4. Gill Arch Condition in the Myrophinae 


Abbreviations as in Table 3. 


By Bo B3 By, Hy Ho H C1-4 Cc Io 13 UP3 UPL 
Benthenchelys cartieri CoS ee ae Oh 050 0 =O 0 =) 
Ahlta egmontis QO os => = @ @ = 0 = = @ @ 0 
Myrophis punctatus* 0 = = @ @ € 0 = @ @ © 0 
M. vafer 0 S = 0 0 (¢ 0 = 0 0 0 0 
M. uropterus Op eee ee ON Olan 0 > @ @ © 0 
M. plumbeus QO ce = @ @ € 0 = @ © =0= 
Pseudomyrophis nimius 0 >= = @ @ € 0 = 7 © @ 0 
P. mteropinna 0) = 5 @ @ € 0 = © @© @ 0 
Neenchelys buttendijkt* R oc Ss & 0 © € 0 > 0 © @ 0 
Sehultzidta johnstonensts = = =] = 0) 05 JC 0 - = 0 -0- 
Sehtsmorhynchus labialis = = @= 70 Oy = 0) = = @ =l0)= 
Muraenichthys chilenstis > =o 5 = @ © € 0 = © © © 0 
M. macropterus = = = = 0) @) 0 - C 0 Oo 0 
M. cooket* 3 oS OO UE 0) “6 © @ 0 
M. gymnotus > = = = @ @ € 0 - =" 20) 30 0 
M. latteaudata - = = = 0 0 C 0 = 0 0 0 0 
M. schultzet = Oe OeeaC 0 = 2 @ =(0= 
M. gymnopterus > = & = @) @ 0 Seas =(0)= 


112 


SERIES 4, V. 41, #1 McCOSKER — EELS 


Table 5. Lateral Line and Cephalic Pore Conditions in Ophichthine Genera 
and Subgenera 


Cephalic pore locations are illustrated in Figure 24. Abbreviations are: 
*, condition of type species unknown; +, present; -, absent; C, lateral 
line ossicles continuous; M, lateral line ossicles moderately separated 
at pores; S, lateral line ossicles separated at pores. 


tp2 pop3 pop4 Lateral Line 
Ossicles 


Ophtchthus = 
Echelus - 
Ophisurus > 
Pisodonophts = 
Quasst remus = 
Ctrrhimuraena* 

Calamuraena 

Jenkinstella 

Echtophis = 
Mystrtophis = 
Aplatophis = 
Brachysomophts* = 
Xyrtas = = = 
Seytalichthys - - - 
Pogonophts - - - 
Evtps - - - 
Leturanus = = = 
Elapsopis - - - 
Phyllophtchthus = = = 
Malvoltophis = = = 
Myrtchthys = = - 


Leet ttt eee t 
' 


Apterichtus <5 + 
Iehthyapus + + 
Cirrteaecula + + 
Stietorhinus + + 
Caecula + + 
Yirrkala +,- - - 
Lamos toma = = = 


Aprognathodon = = - 
Callechelys = = = 
Letharchus = - - 
Leuropharus = = = 
Paraletharchus = = = 


Bascantchthys* = = = 
Alltps = = - 
Phaenomonas - = = 
Ethadophis - = = 
Caralophta = = = 
Leptenchelys = = = 
Dalophts = = - 


= CS es x St Se (2) (eb ise) (op) (5p) QAANNDQNDAANAN FBNFTFTFBEMANNUUNUNUUNVYUNUUNUNNNYN 


PROCEEDINGS 


CALIFORNIA ACADEMY OF SCIENCES 
Table 6. Vertebral Counts of Various Ophichthid Species 


Data were obtained during the present study except as cited. The listing is arranged alpha- 


betically by genus and species within each tribe. Counts include the hypural. SUNY Ufayelti= 
cates number of individuals. 
ee 
Holotype Range Mean N Source 

Benthenchelys cartiert 156-174 168.3 34 Castle, 1972 
Ahlta egmontis 152 l 
Muraentchthys aokt 137 
Muraentchthys australis 152 Castle, 1965 
Muraenichthys breviceps 164 Castle, 1965 
Muraentchthys chilensts 149 =148-153. 150.9 12 £McCosker, 1970 
Muraentchthys cooket 130 Gosline, 195la 
Muraentchthys gymnotus 129-130 129.5 2 
Muraentchthys hattae 154 
Muraenichthys tredalet 126-127. 126.5 2 
Muraenichthys macropterus 127-130 128.2 4 
Muraenichthys schultzet \772 2 
Muraentchthys thompsont 128? 128-133 130.5 2 
Myrophis plumbeus 142-154 Blache, et al., 1970 
Myrophis punctatus 138-145 Eldred, 1966 
Myrophis vafer 146-150 147.8 5 

(holotype of Hesperomyrus fryt) 154? 
Neenchelys buitendijkt 145-148 Mohamed, 1958 
Pseudomyrophis mtcroptnna 174 
Pseudomyrophts ntmtus 212-216 214 2 
Schismorhynchus labialis 136 134-138 136 6 
Schultztdia johnstonensts 145-149 = 151.2 5 


Schultztdia retropinnis 
(holotype of Muraenichthys malaita) 133 


Aprognathodon platyventris 150-155 S275 2 


Callechelys bilinearis 161-163 162 2 
Callechelys elt ffi 155-149-158 154.9 14 
Callechelys ertstigmus 159 154-163-157. 30 
Callechelys galapagensis 172 170-174 172 4 
Calleechelys holochromus 166 
Callechelys Leucopterus 162-165 164 4 Blache and Cadenat, 1971 
Callechelys marmoratus 176-183 179 4 

(holotype of Callechelys guichenoti) 183 
Callechelys melanotaentus 200-205 203 5 
Callechelys muraena 141 41-144 142. 2 
Callechelys nebulosus 158-159 158. 3 
Callechelys perryae 178 Blache and Cadenat, 1971 
Callechelys springert 170 166-170 168 2 
Callechelys striatus 192 | 
Letharehus velifer 139 «135-143 s:«139. 14. McCosker, 1974 
Letharehus rosenblattt 151 144-151 148. 20 McCosker, 1974 
Leuropharus lastops 135 
Paraletharchus opercularis 180 171-180 174. 9 
Paraletharchus pact ficus 166 156-167 160. 15 
Apterichtus ansp 123-132 -  Bohlke, 1968 
Apterichtus caecus 151 ] 
Aptertchtus equatorialis?| 146 1 

(holotype has 53 preanal vertebrae) 
Apterichtus flavicaudus 145-157-149. 5 
Aptertchtus gymnocelus 136 1 
Apterichtus kendalli 137-144 Bohl ke, 1968 
Apterichtus klazingat 140 
Caecula pterygera 126 126-130 128. 8  Bohlke and McCosker, 1975 
Ctrrtecaeecula johnsont 119 ] 
Iehthyapus acuttrostris 133 Blache and Bauchot, 1972 
Iehthyapus ophtoneus 133 321378 lsse 3 
Ichthyapus selachops 137-144 139. 15 


1. The holotype of Caeeula equatorialis Myers and Wade lacks a tail. 


The specimen recorded 


here, ANSP 117436, is from 3015'S, 80°19'W, and was not compared with the type. 


we} 


SERIES 4, V. 41, #1 McCOSKER — EELS © 


Table 6. Continued 
Holotype Range Mean N Source 

Iehthyapus vulturis? 123 

from Palau, Tahiti, and Seychelles N27, Wiis 12 

from Hawaii and Kure 120-124 122. 4 

from Easter Island 130-134 132. 9 
Lamostoma kampent 143 ] 
Lamostoma mindora 144 
Lamostoma orientalis 134-1376 135. 3 
Lamostoma phtltppinensts 153? 
Sttetorhinus potamtus 140 3=—135=142 = 139. 11 = Bohike and McCosker, 1975 
Ytrrkala lLumbricotdes 151-154; 152) 2 

(paratype of Yirrkala chaselingt) 153 
Yirrkala tenuis 153 ] 
Alltps concolor 174 
Bascantchthys cecilae 226 225-226 225. 2 Blache and Cadenat, 197] 
Bascanichthys congoensts 189 189-190 189. 2 Blache and Cadenat, 1971 
Bascantchthys longtssimus 212 - Blache and Cadenat, 197] 
Bascantchthys myerst 215 
Baseantechthys panamensis 181 l 
Bascanichthys paulensts 191 
Bascantchthys tenuts 203 
Basecantchthys teres 181-184 - Blache and Cadenat, 197] 
Caralophta loxochila 139-145 142. 3 
Dalophis tmberbis 152 148-159 152. 14. Blache and Bauchot, 1972 
Ethadophts byrnet 189 
Ethadophis merenda 159 
Leptenchelys vermi formis 163 
Phaenomonas cooperae 270 243-270 256 8 
Phaenomonas pinnata 187 1175-194 1186. 14 
Aplatophis chaultodus 110-111 110. 2 
Brachysomophts henshawt 130 -  Gosline, 195la 
Brachysomophis sauropsis NGS 125 124 4 
Ctrrhtmuraena macgregort 181 -  Gosline, 195la 
Ctrrhimuraena taentopterus 183 1 
Echelus myrus 

from the Mediterranean S55) = Grassi, 1913 

from the eastern tropical Atlantic 149-152 = Blache, et al., 
Echelus pachyrhynchus 157. 149-157 - Blache, 1968 
Echtophis intertinectus 132 132-143 - Blache, 1971 
Echtophis mordax 130 
Echtophtis sp. 132-139 135. 14 
Elapsopts cyeclorhinus 153-160 156. 2 
Evips percinetus 132 
Leturanus semicinetus 162-169 166. 3 

(holotype of Machaerenchelys 

phoenixensis) 170 
Myrichthys bleekeri 196 | 
Myrtchthys colubrinus 197-201 199 2 
Myrichthys maculosus 

from Hawaii and Midway 174-182 178. 16 

from western Pacific 190-199-193. 10 
Myrichthys oculatus 170-171 7). 2 
Myrichthys pardalts 158 151-159 - Blache and Cadenat, 1971 
Myrichthys xystrurus 149-163 9152. 20 
Mystrtophts blastorhinos 142 Blache, 1971 
Mystrtophts crosntert 136-144 140. 57 Blache, 197] 
Mystrtiophts rostellatus 154-158 155. 13. Blache, 1971 
Ophtchthus altiptnnis 173 | 
Ophtchthus callaensis 153 1 
Ophichthus cruentifer 146 l 
Ophtchthus erabo 155 155 2 
Ophichthus frontalis 144-157 149. 7 
Ophtchthus gomest 141 - Jordan and Davis, 1892 


2. Population differences in Ichthyapus vulturis are treated in Randall and McCosker (1974). 
3. An undescribed species ranging from the Gulf of California to Panama. 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Table 6. Continued 


a 


Holotype Range Mean N Source 

Ophichthus maecrochir 144 - Castle, 1965 
Ophichthus ocellatus 134 - Jordan and Davis, 1892 
Ophichthus ophis 

from Brazil 162 l 

from eastern tropical Atlantic 161-170 - Blache, in litt. 
Ophichthus rhyttdodermatotdes 191 1 
Ophichthus triserialis 145-160 152.5 2 
Ophtchthus untsertalis 152? 
Ophichthus urolophus 136 ] 
Ophtchthus zophochir 150-152 = 150.7 3 
Ophisurus serpens 200-208 Blache, in litt. 
Phyllophtchthus xenodontus 169-170 169.5 6 
Pisodonophis boro 171-173-172 2 
Pisodonophits canerivorus 155 I 
Pisodonophis daspilotus 1372) 137=138) 13725 2 
Pogonophts fossatus 166-171 168.5 2 
Quasstremus evtonthas 153 1 
Quasstremus nothochir 139-142 140.8 4 
Seytaltchthys miurus 143-149 = 146 4 


Xyrias revulsus 158 


116 


SERIES 4, V. 41, #1 McCOSKER — EELS 


Table 7. Characteristics of the Ophichthidae and Related Eel Families 


Abbreviations are: F, fused; 0, ossified; R, reduced; S, sutured; +, present; -, 
absent. Data are from this study and various sources, including McAllister (1968), 
Robins and Robins (1970, 1971), and Smith and Castle (1972). 


Frontal Frontal Temporal Gill Arch Branchiostegal 
Condition Commissure Pore Canal Ossification Rays (pairs) 


Ophichthidae F + + 

Ophichthinae 0 15-34 

My rophinae R 20-49 
Congridae F - + 0 8-17 
Muraenesocidae F - +? 0 8-22 
Macrocephenchelyidae F - + 0 8 
Dysommi dae F - - R 9-16 
Xenocongri dae S = ce R 12-21 
Heterenchel yidae S - + 0 11-14 
Muraenidae Ss = = R <16 
Angui | lidae S = + 0 8-14 


ee ——————————————eSSSSSSSSSSSSSSSSSeSSeeSeEeESEeEeEeEeEeEeEEeEEeEeEeEeEeESESESESESESeSESESEeSeses 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES Wala 


Table 8. Morphological and Meristic Characters of the Species of Callechelyini 


Adapted from and expanded upon McCosker and Rosenblatt (1972, Table 5). Abbreviations are 
as follows: 4, rounded mean value; *, from the holotype; +, present; ~-, absent; Ue .condii— 
tion not known. Postanal lengths in thousandths of total length. 


Postanal® Vertebrae Scapula Urohyal Branchiostegal Epihyal Ray 


Length Rays Broadening 
Aprognathodon platyventris 35] 1554 + simple 28 extreme 
Leuropharus lastops 405 135% + ? 32? no? 
Letharchus veltfer 400 149% + 2 32 extreme 
L. rosenblattt 428 151* + simple 34 extreme 
Paraletharchus pact ficus 352 166* + simple 32 yes 
P. opercularis 392 180* + simple 30 yes 
Callechelys nebulosus 408 1594 + simple 29 yes 
C. springert 350 170* + simple 25? yes 
C. btltnearts 364 1624 + forked 27 no 
Gs eiheyage 43h 155% + forked 26 no 
C. galapagensis 4h 172* + forked 27 no 
C. luteus 415 213 + forked 27 no 
C. muraena 385 141% + forked 23? no 
C. marmoratus 345 180° - simple 29 no 
C. ertstigmus 295 158% = simple 31 no 
C. melanotaentus 282 2034 - simple 29 no 
C. striatus 304 192 - simple 26 no 
C. perryae 319 178% - simple 24 no 
C. Leucopterus 453 1644 2 ? ? ? 
C. holoehromus 333 166% - simple ? yes 
C. bttaentatus 385 ? ? ? ? ? 


SERIES 4, V. 41, #1 McCOSKER — EELS 


Table 9. Characteristics of the Species of Callechelyini Used in 
Programs REGROUP and WVGM 


a 


1]. Meristics 


Vertebrae: 130-139; 140-149; 150-159; 160-169; 170-179; 180-189; 190-199; 
200-209; 210-219 

Branchiostegal rays: 22-23; 24-25; 26-27; 28-29; 30-31; 32-33; 34-35 

Supraorbital pores: 3 or 4 


Il. External Morphology 


Postanal length: 275-299; 300-324; 325-349; 350-374; 375-399; 400-424; 
425-449; 450-474 

Anal fin: present or absent 

Gill opening ''pocket'': present or absent 

Underside of snout: grooved, slightly grooved or ungrooved 

Snout: blunt or conical; papillate or smooth 

Anterior nostril: tubular or not tubular 


Ill. Internal Morphology 


DFO: above supraoccipital or above epiotics 

Neurocranium: rounded or depressed 

Intermaxillary teeth: present or absent 

Urohyal: forked, simple and cartilaginous or simple and ossified 

Hyoid: inflexible along CH-EH suture, slightly flexible or well separated 
Hypohyals: present or absent 

Vomerine teeth: present or absent 

Scapula (?): present or absent 

Epihyal rays: broadened basally, moderately broadened or slender 

Third hypobranchial: ossified or cartilaginous 


IV. Coloration 


Mottled or weakly spotted 
Strongly spotted 

Longitudinally banded 

Uniform body (fins may contrast) 


mn 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Table 10. 


Distribution of Certain Ophichthid Genera 


Refer to text for discussion of those genera marked with an asterisk. 


Apterichtus 
Iehthyapus 


Achtrophichthys 
Brachysomophis 
Ctirrt caecula 
Lammos toma 
Muraent chthys* 
Schultatdia 


Echtophts* 


Ethadophts 
Paraletharchus 


Ahlta 
Gorditchthys 


VI. 


Dalophis 


|. Circumtropical 


Bascanichthys 
Myrichthys 
Ophtchthus 


Il. Restricted to Indo-Pacific 


Alltips 
Caecula 
Elapsopts 
Leturanus 
Neenchelys 
Xyrtas 


Ill. Restricted to New World 


Letharchus 
Quasst remus 


IV. Restricted to Eastern Pacific 
Leptenchelys 
Pogonophis 

V. Restricted to Western Atlantic 


Aprognathodon 
Sttetorhinus 


Echelus 


Callechelys 
Myrophis 


Benthenchelys 
Ctirrhimuraena 
Evips 
Malvoltophis 
Sehtsmorhynchus 
Yirrkala 


Pseudomyrophis 


Leuropharus 
Seytalichthys 


Caralophta 


Restricted to Eastern Atlantic and Mediterranean 


Mystrtophts 


119 


120 


SERIES 4, V.41, #1 McCOSKER — EELS 


INDEX TO GENERA AND SPECIES 


(Included are recognized species and valid and invalid genera) 


Acanthenchelys 80,81 
Achirophichthys 65 
acuminatus, Myrichthys 78 
acutirostris, Ichthyapus 67,68 
acutirostris, Muraenichthys 59 
Ahlia 58 

aliculatus, Letharchus 64 
Allips 70 

altipinnis, Ophichthus 80,81 
Anepistomon 82 
anguiformis, Apterichtus 66 
Anguisurus 68 

ansp, Apterichthus 66 
Antobrantia 80 

Aotea 58,59 

apicalis, Ophichthus 80,81 
Aplatophis 74 
Aprognathodon 62 
Apterichtus 59,65,68 
asakusae, Ophichthus 81 
ascensionsis, Ophichthus 81 
ater, Ophichthus 81 
atlanticus, Brachysomophis 74 
australis, Muraenichthys 58,59 
australis, Myrophis 60 
Bascanichthys 70,78 
bascanoides, Bascanichthys 71 
Benthenchelys 57,85 
bilinearis, Callechelys 63 
bitaeniatus, Callechelys 63 
blastorhinos, Echiophis 77,79 
bleekeri, Myrichthys 78 
bonaparti, Ophichthus 80,81 
boro, Pisodonophis 82 
boulengeri, Dalophis 72 
Brachycheirophis 83 
Brachysomophis 65,74,75,79 
Branderius 65 

breviceps, Muraenichthys 59 
buitendijki, Neenchelys 60 
byrnei, Ethadophis 72 
Caecilia 65 

Caecula 63,65,66,67,69,70,72,85 
caecus, Apterichtus 65,66 
Calamuraena 75 

calamus, Cirrhimuraena 75 
Callechelys 62,72 

callaensis, Ophichthus 81 


cancrivorus, Pisodonophis 82,83 


Caralophia 71 
cartieri, Benthenchelys 57 
ceciliae, Bascanichthys 71 


celebicus, Ophichthus 81 
Centrurophis 79,81 
cephalopeltis, Dalophis 71,72 
cephalozona, Ophichthus 79,80,81 
chauliodus,Aplatophis 74 
cheilopogon, Cirrhimuraena 75 
cheni, Myrophis 60 

chilensis, Muraenichthys 59 
chinensis, Cirrhimuraena 75 
Chlevastes 78 

Cirrhimuraena 75,85 
cirrocheilos, Brachysomophis 74,75 
Cirricaecula 59,67,68 

cliffi, Callechelys 63 
Coecilophis 80,81 

Cogrus 79,81 

colubrinus, Myrichthys 78 
concolor, Allips 70 
congoensis, Bascanichthys 71 
cookei, Muraenichthys 59 
cooperae, Phaenomonas 73 
copelandi, Pisodonophis 82 
crocodilinus, Brachysomophis 65,74 
crosnieri, Mystriophis 79 
Crotalopsis 76,77 

cruentifer, Ophichthus 80,81,82 
Cryptopterenchelys 80 
Cryptopterus 80,81 
Cryptopterygium 62,63 
Cyclophichthys 77 
cyclorhinus, Elapsopis 77 
cylindricus, Bascanichthys 71 
cylindroideus, Myrophis 59,60 
Dalophis 62,68,71 

daspilotus, Pisodonophis 82,83 
derbeyensis, Ophichthus 81 
devisi, Muraenichthys 59 
dromicus, Pisodonophis 82 
Echelus 75,76,81,85 

Echiophis 76,77,79 

Echiopsis 76 

egmontis, Ahlia 58 

Elapsopis 77 

epinepheli, Apterichtus 66 
episcopus, Ophichthus 81 
equatorialis, Apterichtus 66 
erabo, Ophichthus 81 
eristigmus, Callechelys 63 
Ethadophis 72 

evermanni, Ophichthus 81 
evionthas, Quassiremus 83,84 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 


Evips 77 

filaria, Bascanichthys 71 
flavicaudus, Apterichtus 68 
foresti, Apterichtus 65,66 
fossatus, Pogonophis 83 
frio, Myrophis 60 

frontalis, Ophichthus 81 
fusca, Yirrkala 69 
galapagensis, Callechelys 63 
garretti, Ophichthus 81 
Gisenchelys 80,81 
gjellerupi, Yirrkala 69 
godeffroyi, Muraenichthys 59 
gomesi, Ophichthus 80,81 
Gordiichthys 63,72 

gracilis, Apterichtus 65 
grandoculis, Ophichthus 81 
gymnocelus, Apterichtus 66 


gymnopterus, Muraenichthys 58,59 


gymnotus, Muraenichthys 59 
hattae, Muraenichthys 59 
Hemerorhinus 67,70 

henshawi, Brachysomophis 75 
Herpetoichthys 80 
Hesperomyrus 59 

heyningi, Hemerorhinus 67 
hijala, Pisodonophis 82 
hoevenii, Pisodonophis 82 
holochromus, Callechelys 62,63 
hypselopterus, Pisodonophis 82 
Ichthyapus 59,66,67,68 
imberbis, Dalophis 71,72 
inhacae, Cirrhimuraena 75 
Innominado 79,81 

intertinctus, Echiophis 76,77 
iredalei, Muraenichthys 59 
irretitus, Gordiichthys 72 
Jenkinsiella 75,85 

johnsoni, Cirricaecula 64 
johnstonensis, Schultzidia 61,62 
kampeni, Lamnostoma 65 
kaupi, Yirrkala 69,70 

kendalli, Apterichtus 65 

kirkii, Bascanichthys 71 
klazingai, Apterichtus 66 
labialis, Schismorhynchus 61 
Lamnostoma 65,67,68,69,85 
lasiops, Leuropharus 64 
laticaudata, Muraenichthys 58,59 
Leiuranus 77 

Leptenchelys 72,73,85 
Leptognathus 82 
Leptorhinophis 80 
Leptorhynchus 82 


lepturus, Myrophis 60 
Letharchus 63,64 

leucopterus, Callechelys 63 
Leuropharus 64 

limkouensis, Ophichthus 81 
longissimus, Bascanichthys 71 
loxochila, Caralophia 71 
lumbricoides, Yirrkala 69,70 
luteus, Callechelys 63 
macgregori, Cirrhimuraena 75 
Machaerenchelys 77 
macrochir, Ophichthus 81 
macrodon, Yirrkala 69 
Macrodonophis 76 

macrops, Ophichthus 81 
macropterus, Muraenichthys 59 
macrorhynchus, Ophisurus 82 
macrostomus, Muraenichthys 59 
macrurus, Phyllophichthus 82 
maculata, Yirrkala 69 
maculatus, Ophichthus 79,81 
maculosus, Myrichthys 78,79 
madagascariensis, Ophichthus 81 
magnioculis, Ophichthus 80,81 
Malvoliophis 77,78,85 
manilensis, Ophichthus 81 
marginatus, Ophichthus 81 
marmoratus, Callechelys 62,63 
melanochir, Ophichthus 81 
melanoporus, Ophichthus 81 
melanotaenius, Callechelys 63 
merenda, Ethadophis 72 
Microdonophis 75,80,81,85 


micropinna, Pseudomyrophis 60,61 


Microrhynchus 65-66 
microtretus, Neenchelys 60 
mindora, Lamnostoma 69 
misolensis, Yirrkala 69,70 
miurus, Scytalichthys 84 
moluccensis, Yirrkala 69 
monodi, Apterichtus 66 
mordax, Echiophis 76,77 
moseri, Apterichtus 
multidentatus, Dalophis 72 
multiserialis, Ophichthus 81,82 
muraena, Callechelys 63,65 
Muraenichthys 58,59,61,62,73 
Muraenophis 80 
Muraenopsis 80,81 

myersi, Bascanichthys 71 
Myrichthys 78,79 

Myrophis 57,58,59 
Myropterura 58 

Myrus 75,76 


121 


122 


myrus, Echelus 75 
Mystriophis 77,79,84 
natalensis, Yirrkala 69 
nebulosus, Callechelys 63 
Neenchelys 60,85 

nimius, Pseudomyrophis 60,61 
nothochir, Quassiremus 84 
obtusirostris, Dalophis 72 
oculatus, Myrichthys 78 
oliveri, Cirrhimuraena 75 
omanensis, Ichthyapus 68 
Omochelys 80,81,82,85 
opercularis, Paraletharchus 64 


Ophichthus 69,73,75,76,79,80,81,82,83,85 


ophioneus, Ichthyapus 67,68 
Ophis 79,80,81 

Ophisuraphis 65 

Ophisurapus 65 

Ophisurus 74,76,77,78,79,80,81,83 
Ophithorax 79,81 

opici, Hemerorhinus 67 
orientalis, Lamnostoma 68,69 
Oxydontichthys 80 
pachyrhynchus, Echelus 76 
pacificus, Ophichthus 81 
pacificus, Paraletharchus 64 
panamensis, Bascanichthys 71 
Pantonora 69,70 
Paraletharchus 64 

Paramyrus 59 

pardalis, Myrichthys 79 

parilis, Ophichthus 81 
paucidens, Cirrhimuraena 75 
paulensis, Bascanichthys 71 
Pelia 71,72 

percinctus, Evips 77 

perryae, Callechelys 63 
Phaenomonas 61,73 
philippinensis, Muraenichthys 59 
Phyllophichthus 78,82 

pictum, Lamnostoma 67,68 
pinguis, Malvoliophis 78 
pinnata, Phaenomonas 73 
Pisodonophis 79,80,81,82,83,85 
platyrhynchus, Myrophis 60 
platyventris, Aprognathodon 62 
playfairi, Cirrhimuraena 75 
plumbeus, Myrophis 59,60 
Poecilocephalus 79,81 
Pogonophis 83 
polyophthalmus, Ophichthus 81 
porphyreus, Mystriophis 79 
potamius, Stictorhinus 69 
productus, Quassiremus 84 


SERIES 4, V. 41, #1 McCOSKER — EELS 


Pseudomyrophis 60,61 
Pterurus 71 

pterygera, Caecula 66 
punctatus, Myrophis 59,60 
puncticeps, Ophichthus 80,81 
punctifer, Echiophis 76,77 
pusillus, Bascanichthys 71 
Quassiremus 83 

reguis, Ophichthus 81 
remiger, Ophichthus 81 
retifer, Ophichthus 81 
retropinna, Schultzidia 81 
retropinnis, Ophichthus 62 
revulsus, Xyrias 84 
Rhinenchelys 67,68 
rosenblatti, Letharchus 64 
roseus, Ophichthus 81 
rostellatus, Mystriophis 79 
rufus, Ophichthus 81 
rutidoderma, Ophichthus 81 
rutidodermatoides, Ophichthus 81 
sauropsis, Brachysomophis 75 
Schismorhynchus 59,61 
schultzei, Muraenichthys 59 
Schultzidia 59,61 
Scolecenchelys 58,59 
Scytalichthys 84 

Scytallurus 71 

Scytalophis 80,81 

selachops, Ichthyapus 68 
semicinctus, Leiuranus 77,78 
semicinctus, Pisodonophis 82 
serpens, Ophisurus 81,82 
sibogae, Muraenichthys 59 
Sphagebranchus 65,66,67,68,69,70,71,72,77 
spinicauda, Ophichthus 80,81 
springeri, Callechelys 63,72 
stenopterus, Ophichthus 81 
Stethopterus 77 

Stictorhinus 69 

striatus, Callechelys 63 
Syletophis 80 

Syletor 80 

taeniopterus, Cirrhimuraena 75 
taylori, Lamnostoma 69 
tenuis, Bascanichthys 71,73 
tenuis, Yirrkala 69 

teres, Bascanichthys 70,71 
thompsoni, Muraenichthys 59 
triserialis, Ophichthus 81 
Typhlotes 65 

typus, Achirophichthys 65 
unicolor, Ophichthus 81 
Uranichthys 80,81 


PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES 123 


urolophus, Ophichthus 81 woosuitingi, Ophichthus 81 
uropterus, Myrophis 60 xenodontus, Phyllophichthus 82 
vafer, Myrophis 59,60 xorae, Muraenichthys 59 
velifer, Letharchus 63,64 Xyrias 84 

Verma 65,66 xystrurus, Myrichthys 78,79 
vermiformis, Leptenchelys 72 Yirrkala 67,69 

vermiformis, Muraenichthys 59 Zonophichthus 80,81,85 
versicolor, Elapsopis 77 zophistus, Pisodonophis 82 


vulturis, Ichthyapus 68 zophochir, Ophichthus 75,80,81 


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NH 
PROCEEDINGS 
OF THE 
CALIFORNIA ACADEMY OF SCIENCES 
FOURTH SERIES 
Vol. XLI, No. 2, pp. 125-160, 16 figs., 11 tables April 15, 1977 


STUDIES ON EASTERN PACIFIC SAND STARGAZERS 
(PISCES: DACTYLOSCOPIDAE) 4. G/LLELLUS, SINDOSCOPUS NEW GENUS, 
AND HETERISTIUS WITH DESCRIPTION OF NEW SPECIES 


By 


C. E. Dawson 


Gulf Coast Research Laboratory Museum 
Ocean Springs, Mississippi, 39564 


ABSTRACT: The dactyloscopid genera Gillellus and Heteristius are diagnosed and described 
and a new genus, Sindoscopus, is proposed for the Chilean endemic, Gillel/us australis Fowler 
and Bean. A key is provided for all Pacific genera and the seven species treated here. Gillellus 
includes one species, G. searcheri n. sp. (Islas Tres Marias and Nayarit, Mexico and Costa 
_ Rica to Panama), with some paired infraorbital pores. There are four more closely related 
species without paired pores. The latter group includes the generic type, G. semicinctus Gil- 
bert (Isla Guadalupe, Baja California peninsula and western Gulf of California, mainland 
shores from Nayarit, Mexico to Colombia, and the Revillagigedos and Galapagos Islands). 
Gillellus arenicola Gilbert occurs in the vicinity of Cape San Lucas, fails to enter the Gulf 
of California, and has been collected off mainland coasts of Colima and Oaxaca, Mexico. Gill- 
ellus ornatus Gilbert is apparently restricted to the Gulf of California where all but one speci- 
men has been taken along its western shores. Gillellus chathamensis n. sp. is an insular en- 
demic known only from Cocos Island, Costa Rica. The monotypic Heteristius occurs along the 
southern portion of Baja California. It is not recorded from the Gulf of California. This species 
also appears off Mexican mainland shores (Isla Tres Marias, Nayarit to Oaxaca) and is found 
off Costa Rica, Panama and Ecuador (to about 00°27’N). Heteristius jalisconis Myers and Wade, 
type species of Heteristius, is considered a junior synonym of Dactyloscopus cinctus Osburn 
and Nichols. Sindoscopus, also monotypic, is known only from the Chilean coast (ca. 23°S- 
33°S). All treated species are illustrated and distribution maps are provided for species of 
Gillellus and Heteristius. This represents the final part of a review of Pacific dactyloscopids 
wherein a total of 7 genera and 29 species or subspecies are recognized. 


CALIFORNIA ACADEMY OF SCIENCES/GOLDEN GATE PARK/SAN FRANCISCO 


126 


INTRODUCTION 


This report, the last of a series reviewing 
Pacific dactyloscopids, treats known populations 
of Gillellus Gilbert and Heteristius Myers and 
Wade, and a new genus, Sindoscopus, is pro- 
posed for the Chilean endemic, Gillellus australis 
Fowler and Bean. 


As with other Pacific sand stargazers, these 
forms have never been adequately treated sys- 
tematically, existing illustrations are few and in- 
accurate, and identification has been difficult or 
impossible. Material from the Revillagigedos Is., 
mainland coast of South America and the Gala- 
pagos Is. is minimal and further study is needed 
in these areas. Nevertheless, present data Clarify 
the systematic status of all known museum hold- 
ings and establish a basis for identification of 
future collections. 


METHODS AND MATERIALS 


Counts and measurements, recorded in milli- 
meters (mm), follow methods of Dawson (1969, 
1974, 1975). Standard length (SL) is used through- 
out; counts of meristic characters are total 
counts; preopercular canal pores are enumerated 
in the 1st (anterior) primary canal (Fig. 1). Counts 
of lateral-line scales were routinely made on the 
right side, but left side counts were substituted 
where right side squamation was incomplete. The 
posteriormost short spiniform caudal elements 
may occasionally be irregularly segmented in 
Gillellus and Sindoscopus, but these are omitted 
from counts of principal rays which include only 
the long, well-developed, segmented elements. 
Branching of principal segmented caudal rays is 
indicated by the formula: number of upper 
simple rays + number of branched rays + 
simple lower rays. Measurements are given for 
undamaged primary types except for Sindoscopus 
australis where a specimen of similar length has 
been substituted for the damaged holotype. Ob- 
servations on dentition and general osteological 
features are from one or more cleared and 
stained specimens of each species; vertebral 
counts are from radiographs. Materials examined 
include virtually all known museum holdings, but 
meristic data were not taken on all specimens. 

Abbreviations for repositories of examined 
material are: AMNH - American Museum of Nat- 
ural History; ANSP - Academy of Natural Sciences 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


of Philadelphia; BC - Institute of Animal Resource 
Biology, University of British Columbia; CAS - 
California Academy of Sciences; CAS-SU - Stan- 
ford University specimens now housed at CAS; 
GCRL - Gulf Coast Research Laboratory Museum; 
LACM - Los Angeles County Museum of Natural 
History; MNHN- Museo Nacional de _ Historia 
Natural, Santiago, Chile; SIO - Scripps Institution 
of Oceanography; UCLA- University of Cali- 
fornia, Los Angeles; UCR - Universidad de Costa 
Rica, San José; USNM- National Museum of 
Natural History, Smithsonian Institution. 


KEY TO GENERA AND SELECTED SPECIES OF 
PACIFIC DACTYLOSCOPIDS 


1a. Dorsal-fin origin on nape -_...---.---.--------- 2 
1b. Dorsal-fin origin behind nape, near verti- 
cal from: anal-fin origin —- Se 10 


2a. Dorsal fin usually discontinuous; princi- 
pal preopercular canals 3 or more than 6 


2b. Dorsal fin continuous; principal preop- 
erqular canals: 4 22 eee 
....Sindoscopus australis (Fowler and Bean) 


3a. Anterior dorsal-fin elements a series of 
free spines; eyes on protrusile stalks __ 
__Dactyloscopus Gill (see Dawson, 1975) 

3b. Anterior dorsal-fin elements united to 
form a distinct 3 or 2-spined finlet; eyes 
protruding but not stalked -__.......---... - 


4a. Principal segmented caudal rays 10; pec- 
toral rays modally 12 or 13; upper lip 
without fimbriae; venter always naked... 5 

4b. Principal segmented caudal rays more 
than 10; pectoral rays modally 14; fim- 
briae present on upper and lower lips; 
venter at least partly scaled in late juv- 
eniles and adults __............. ees 9 


5a. Dorsal spines total 11-15; 18-29 scales in 
straight portion of lateral line; infraorbi- 
tal canal pores in single series ---........... 6 
5b. Dorsal spines total 20-23; 9-13 scales in 
straight lateral line; some paired pores in 
infraorbital canal _... ee 
ede ete) et eT Gillellus searcheri sp. nov. 


6a. Anterior dorsal finlet 3-spined -.-.-.....-..- 7 
6b. Anterior dorsal finlet 2-spined ---........... 8 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


7a. Pectoral rays usually 12; segmented anal 
rays 30-35 (usually 32-34); 18-23 (usually 
19-21) scales in straight lateral line __...... 
“oe he me Gillellus semicinctus Gilbert 

7b. Pectoral rays usually 13; segmented anal 
rays 33-36 (usually 34-35); 23-25 (usually 
24) scales in straight lateral line —......_. 
2 Gillellus chathamensis sp. nov. 


8a. Tip of lower jaw neither distinctly fleshy 
nor strongly protruding anteriad; 22-26 
scales in straight lateral line; lateral line 
deflects between verticals from dorsal-fin 
elements 18-22 _...Gillellus ornatus Gilbert 

8b. Tip of lower jaw fleshy and strongly 
protruding in undamaged fish; 27-29 
scales in straight lateral line; lateral line 
deflects between verticals from dorsal- 
FIMPCICMMICE ES P4502 ooccaceoen oe ee 


9a. Principal segmented caudal rays 11, 
some branched in adults; dorsal finlet 
3-spined; narrow band of scales across 
venter and scales on pectoral-fin base in 
late juveniles and adults; principal pre- 
Spcetlanwednals: We. a 
....Platygillellus Dawson (see Dawson, 1974) 

9b. Principal segmented caudal rays 12, un- 
branched; dorsal finlet 2-spined; venter 
fully scaled; pectoral-fin base naked in 
late juveniles and adults; principal pre- 
opercular canals more than 6 _............ 
...._Heteristius cinctus (Osburn and Nichols) 


10a. Head truncate to broadly rounded in 
front; upper lip fimbriae 12-21, usually 
more than 13; infraorbital canal pores 
TOTS! TT ET LC) ee ea ace ce ea 
ee his Dactylagnus Gill (see Dawson, 1976) 

10b. Head conically pointed in front; upper 
lip fimbriae 4-13, usually less than 13; 
infraorbital canal pores less than 10, typi- 
eal | Ve GaN ge et ee ee ere 
————— Myxodagnus Gill (see Dawson, 1976) 


SYSTEMATIC DESCRIPTIONS 


Gillellus Gilbert, 1890: 98 (type-species by original designa- 


) 
Genus Gillellus Gilbert 
| tion, Gillellus semicinctus Gilbert, 1890). 


DIAGNOSIS: With separate 2- or 3-spined dorsal 
finlet originating on nape, with or without iso- 


| 


127 


lated spines between finlet and remainder of 
dorsal fin; eyes not stalked, usually with a minute 
flap or papilla; pectoral fin-rays modally 12 or 
13; principal segmented caudal rays 10, some 
branched in late juveniles and adults; lower lip 
with up to 7 fleshy fimbriae, upper lip entire; 
head, pectoral-fin base and venter naked; princi- 
pal preopercular canals 3, unbranched; without 
predorsal bones (interneurals). 


DESCRIPTION: Body rather slender, broader and 
deeper in front, laterally compressed and narrow- 
ing caudad; head moderately large, rounded dor- 
sad, broadest near posterior margin of pre- 
opercle, somewhat narrowed in front; lower jaw 
slightly to strongly protruding, rounded in dorsal 
profile; upper lip entire, smoothly rounded and 
devoid of broad emarginations or fimbriae; lower 
lip with up to 7 (usually 4 or 6) unbranched 
fleshy papilla-like fimbriae (Fig. 5); eyes dorso- 
lateral, protruding but not on long protrusile 
stalks, usually with a flap or papilla; anterior 
naris tubiform, the aperture simple or with valve- 
like emargination, located on preorbital; poster- 
ior naris, a simple pore with slightly elevated rim, 
located behind tubiform naris and adjacent to 
anterior rim of orbit; principal preopercular can- 
als 3, unbranched, and typically with but a single 
pore in the 1st (anterior) canal; infraorbital canal 
pores in single series or paired posteriad (Fig. 1); 
opercle with a fringe of up to 12 short, usually 
unbranched fimbriae overlying upper posterior 
margin, the frequency ontogenetic (Fig. 7); 
opercles broadly expanded, the membranous 
margins cover pectoral- and pelvic-fin bases and 
overlap on underside of head. 

Dorsal-fin origin on nape, in advance of upper 
opercular angle; dorsal fin with an isolated 2- 
or 3-spined anterior finlet, 1st and 2nd spines 
more closely spaced than 2nd and 3rd; when 2- 
spined, finlet is followed by one or two isolated 
spines; remainder of spinous dorsal continuous 
with a series of segmented rays behind, the fin 
slightly to deeply emarginate between; anal 
spines 2, the 1st about a third shorter than 2nd 
which is similarly shorter than 1st segmented 
ray; anterior anal-fin elements of mature (trans- 
formed) males modified, the spines angled cau- 
dad, 1st and 2nd segmented rays angled forward, 
modified segmented rays not appreciably thick- 
ened or fleshy but membranes stretched or 
elongated between. Posteriormost dorsal and 
anal rays usually free, occasionally bound to 
caudal peduncle by short membranes; last anal 
ray reaches to or beyond vertical through rear of 


128 


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PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


Configuration of infraorbital pores and preopercular canals. Top: Gillellus semi- 


cinctus (left), G. searcheri (right). Bottom: Sindoscopus australis (left), Heteristius cinctus 
(right). Principal preopercular canals numbered 1-4; note complete abdominal squamation 


in Heteristius. 


hypural, last dorsal ray usually shorter; caudal 
peduncle entire, without notch in ventral margin 
(Fig. 2); caudal fin rounded, upper and lower- 
most rays typically simple, lower ray usually the 
shorter, middle rays simple in early juveniles but 
branch ontogenetically in larger fish (Table 9). 
Pectoral fin-rays 11-14, most frequently 12 or 
13; pectoral fin narrowly rounded to pointed, 
5th-7th from dorsalmost rays the longest, rays 
gradually shorter ventrad but lower rays not ap- 
preciably swollen or thickened; tip of straight- 
ened and adpressed pectoral fin may reach to 


or slightly beyond descending portion of lateral- 
line arch, usually falls short of descending arch; 
pectoral fin somewhat elongate and cupped or 
outwardly convex in transformed males; pelvic 
fin with a short spine closely bound to 1st of 
three segmented rays, the tips free with emargin- 
ate membranes between; longest pelvic ray may 
reach anal-fin origin but usually falls short of 
anus; all fin-rays simple except for branched 
caudal rays of late juveniles and adults. 

Lateral line continuous, originates just above 
and in advance of upper opercular angle (near 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


vertical from 1st dorsal spine), arches upward to 
follow near dorsal-fin base, deflects (between 
14th-32nd _ dorsal-fin supports) and continues 
midlaterally to terminate as the penultimate scale 
on caudal-fin base; canal of straight lateral-line 
scales parallel to longitudinal body axis, that 
of last scale not strongly angled ventrad 
(Fig. 2). Scales of upper portion of arched lateral 
line persistent, outwardly convex and more or 
less pointed posteriad; scales of descending arch 
and straight lateral line thin and often deciduous, 
rounded in outline, the canal straight and with- 
out a distinct terminal branch anteriad, often 
with a minute dorsal pore on anterior third of 
canal. Squamation variable above |lateral-line 
arch, naked in early juveniles and some adults, 
fully or partially scaled in others; head, predor- 
sum, venter, sides below line between upper 
angle of pectoral axil and 1st to 3rd anal rays, 
and all fins (except caudal base) naked; 5-9 longi- 
tudinal scale rows cross vertical near origin of 
straight lateral line. Premaxillaries somewhat pro- 
tractile, the pedicels reach middle or posterior 
third of orbit but fall well short of posterior mar- 
gin; jaw teeth small, none distinctly enlarged, 
conical and more or less bluntly pointed, in 2-3 
rows near symphyses but mainly uniserial pos- 
teriad; vomer and palatines edentate; tongue 
rounded in front; basihyal slender, barely spatu- 
late near tip. Three complete gill arches plus a 
hemibranch; no gill rakers on outer margin of 
1st arch, often with a few rudimentary rakers 
that barely pick up stain on inner margin; 
pseudobranchiae present; branchiostegals 6; 
without predorsal bones (interneurals); proximal 
pterygiophore of 1st dorsal spine usually inserted 
in advance of 1st neural spine (Fig. 3); abdomi- 
nal vertebrae 10-12, modally 11 (from  radio- 
graphs); caudal fin with 2-4 simple spiniform ele- 
ments above and below, upper and lower hypur- 
als each bear 5 segmented rays. 


Recognized species marked (in alcohol) with 
6-10 principal dark saddle-like bands crossing 
dorsum between dorsal-fin origin and caudal- 
fin base, these often supplemented with second- 
ary bands between and seldom extend ventrad 
of lateral midline; body and fins elsewhere plain, 


_ flecked or spotted with brown; ground color 


pale to brownish. 


REMARKS: Gillellus is similar to Platygillellus 
Dawson, Leurochilus Bohlke and_ Heteristius 
Myers and Wade in possessing an isolated (or 


_ semi-isolated) dorsal finlet inserted on the nape 


129 


Fig. 2. Posterior body and fins, illustrating 
straight lateral-line scales and adult development 
of segmented caudal rays, in Gillellus (top), Sin- 
doscopus (middle) and Heteristius (bottom). 


(Table 1). In this feature it is immediately separ- 
able from Dactyloscopus Gill (continuous dorsal 
fin usually preceded by isolated spines) as well as 
from Dactylagnus Gill and Myxodagnus Gill where- 
in dorsal fin is continuous and originates well be- 
hind nape; these genera also have well devel- 
oped upper lip fimbriae (absent in Gillellus). 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


130 


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DAWSON: EASTERN PACIFIC SAND STARGAZERS 


TABEE 2: 


Gillellus, Sindoscopus and Heteristius. 


Genus and species 


N 11 12 


18) 


Dorsal spines 
14 15 16 17 18 


Frequency distributions of dorsal spines, segmented dorsal rays and total dorsal-fin supports in Pacific 


21 


131 


species of 


22 23 


Gillellus 
semicinctus 
chathamensis 
ornatus 
arenicola 
searcheri 


Sindoscopus 
australis 

Heteristius 
cinctus 


190 


30% © 


87* 


39 


48 3 


16 


19 


Segmented dorsal rays 
21 22 23 24 25 26 27 


32 33 


Gillellus 


semicinctus 
chathamensis 
ornatus 
arenicola 
searcheri 


Sindoscopus 
australis 

Heteristius 
cinctus 


ZO 93" 74 3 


54 


40 


4 42 43 4445 


46 


47 


48 


Gillellus 
semicinctus 
chathamensis 
ornatus 
arenicola 
searcheri 


Sindoscopus 
australis 

Heteristius 
cinctus 


46* 127 


58 


16 


12° 


Geyer she! 4 


(0) 227/ 


*Primary type. 


132 


Flgee 3: 


(left), G. arenicola (right). Middle: G. ornatus (left), G. searcheri (right). Bottom: Sindosopus 
australis (left), Heteristius cinctus (right). PP - proximal pterygiophore; DSI - 1st dorsal spine. 


Platygillellus has 11 principal segmented caudal 
rays and modally 14 pectoral rays (caudal rays 
10, modally 12 or 13 pectoral rays in Gillellus), 
fimbriae occur on both lips, and late juveniles 
and adults have scales on venter and _ pectoral- 
fin base (naked in Gillellus). The monotypic At- 
lantic genus Leurochilus differs from Gillellus in 
having 11 rather than 10 principal caudal rays, 
longer premaxillary pedicels (reach posterior 
edge of orbits), and in the complete absence of 
labial fimbriae. For comparison with Sindoscopus 


and Heteristius see Key and descriptions of these 
genera below. 


Gillellus is represented in Pacific waters by four 
closely related species and one somewhat aber- 
rant form (G. searcheri). This is apparently a 
stenohaline genus and no collections are re- 
corded from low salinity habitats. Maximum size 
is about 50 mm SL. Two species have been 
dredged in 137 m. The genus occurs from Isla 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


Seat 
v 

=< (6) 
n 


LMU 


WEL 


Pp 


Anterior abdominal vertebrae and dorsal-fin supports. Top: Gillellus semicinctus 


Guadalupe, Baja California Norte, Mexico (ca. 
28°53’N) south to the mainland coast of Ecuador 
and the Galapagos Is. (Archipiélago de Colon). 


Gillellus semicinctus Gilbert 
(Figure 4.) 


Gillellus semicinctus Gilbert, 1890: 98 (original description, 
Gulf of California [and Atlantic). 


DIAGNOSIS: Dorsal finlet 3-spined, without iso- 
lated spines between finlet and remainder of 
dorsal fin; pectoral fin-rays modally 12; seg- 
mented anal rays modally 33; 18-23 (usually 19- 
21) scales in straight lateral line; eye flap usually 


present; without paired pores in_ infraorbital 
canal. 


DESCRIPTION: Measurements (mm) of 27.8-mm- 
SL male lectotype follow: depth of caudal pe- 
duncle 1.2; body depth 3.6; predorsal length 6.0; 
preanal length 9.1; pelvic to anal distance 3.6; 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


rs : oe , 
ie . ad > 
a 4 te 


Se ae 


Fig. 4. Gillellus semicinctus Gilbert. Top and middle: Strongly pigmented with indica- 
tions of secondary bands between primary bars crossing dorsum (GCRL 13244, 36 mm SL, 
female). Bottom: Pale pattern lacking secondary bands (CAS 31810, 32 mm SL, female). 


head length 6.6; maxillary to upper opercular 
angle 6.1; maxillary to upper preopercular angle 
4.4: length of anterior naris 0.2; diameter of 
bony orbit 1.4; postorbital length 4.0; snout 
length 1.2; length of maxillary 2.1. See Tables 2-8 
for meristic data. 

Lower jaw slightly protruding; lower lip typi- 
cally with 4 fleshy fimbriae or papilla-like emar- 
ginations, the inner pair the larger (Fig. 5); pre- 


orbital short, rather steeply inclined; tubiform 
anterior naris located close to anterior rim of 
preorbital; eye typically with a minute hyaline 
flap or papilla. Height of dorsal finlet equal to 
or slightly greater than height of remaining spin- 
ous dorsal fin, somewhat less (15-30%) than 
longest segmented dorsal rays (Fig. 6); last dorsal 
spine about 15-20% shorter than 1st segmented 
ray. Pectoral fin-rays 12 in 86% of 437 counts, 3 


PROG. CALIF. ACAD, SGI_, 47H SER, VOL. 41, NO: 2 


Fig. 5. Delineations of anterodorsal aspect of head. Top: Gillellus semicinctus (left), 
G. searcheri (right). Middle: G. arenicola. Bottom: Sindoscopus australis (left), Heteristius 
cinctus (right). Sensory canals and pores omitted. 


fish had 11 rays in each fin, whereas 9 had bi- 
lateral counts of 13; pectoral fin not greatly 
elongate or expanded in transformed males, tip 
of adpressed fin reaches vertical between dorsal- 
fin elements 16 and 19, between 14 and 19 in 


young and females; caudal fin formula modally 
1 + 7 + 2 in examined material (Table 9); op- 
ercular fimbriae 4-12 (Fig. 7; Table 10); infra- 
orbital canal pores usually 7, none paired (Fig. 1). 

Arched portion of lateral line deflects between 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


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Fig. 6. Lateral and dorsal views illustrating lateral-line configuration and squamation 
above lateral-line arch in Gillellus semicinctus (top), G. arenicola (middle) and G. ornatus 


(bottom). 


verticals from dorsal-fin elements 17 and 24, usu- 
ally between 20 and 23; squamation variable 
above lateral-line arch, naked in early juveniles 
and some adults, usually with one or two scale 
rows anteriad above 2nd or 3rd lateral-line scale, 
narrowing to a single row for a distance of some 
10-11 scales and replaced posteriad by the dor- 
salmost portions of 5-6 lateral-line scales in ad- 
vance of lateral-line deflection; anteriormost 
scales often embedded, occasionally isolated; 
area between nape, origin of lateral lines and 
base of 1st or 2nd dorsal spine naked (Fig. 6); 
8-9 longitudinal scale rows cross vertical near 
origin of straight lateral line; scales present in 
arched portion of lateral line in 13-mm fish, 
squamation may be complete in fish about 20 
mm SL. 

Lower jaw with a patch of teeth 2-3 rows 
broad and about 4-5 rows long near symphysis, 
uniserial posteriad; anterior teeth similar in up- 
per jaw but distinctly biserial behind; pseudo- 
branchiae 4-5 (7 specimens examined); proximal 
pterygiophore of 1st dorsal spine broad (Fig. 3). 


Color in alcohol. —Usually with an indistinct 
bar crossing interorbital and continued below 


eye to near posterior angle of gape; often with 
flecks of brown on snout, underside of lower 
jaw, upper portion of preopercle and predor- 
sum; lips and labial fimbriae usually pale. Dor- 
sum crossed by 6-8 (usually 7) principal saddle- 
like bars between nape and caudal-fin base; 
these primary bars equal to or slightly narrower 
than pale interspace, the 1st terminates ventrad 
near middle of opercle, last covers most or all 
of caudal base, whereas remainder terminate 
near lateral midline; well pigmented (usually 
large) specimens often with short secondary bars 
crossing dorsum between primary markings, nu- 
merous brown flecks on otherwise pale inter- 
spaces and two or more primary bars may be 
united ventrad by a narrow brown line; sides 
below midline mainly pale but occasionally with 
scattered brown flecks; pectoral fins usually pale 
but upper interradial membranes may be lightly 
streaked with brown, other fins typically im- 
maculate. Shape and pigmentation of lateral por- 
tions of primary bars (behind nape) variable; 
within a single sample, bars may be narrowed 
between lateral midline and dorsal-fin base, rec- 
tilinear, or narrowed below and somewhat tri- 


136 


SG. semicinctus 
SSS G. chathamesis 


OPERCULAR FIMBRIAE 
OPERCULAR FIMBRIAE 


O 20 60 100 0 20 
STANDARD LENGTH (MM) 


G. arenicola 
G. searcheri 
G. ornatus 


STANDARD LENGTH (MM) 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


S australis 


SS SS SS H. cinctus 


OPERCULAR FIMBRIAE 


(0) 
60 100 O 20 60 100 
STANDARD LENGTH (MM) 


Fig. 7. Regressions of frequencies of opercular fimbriae in Pacific species of Gillellus, 
Sindoscopus and Heteristius. See Table 10 for supporting data. 


angular; bars may have evenly distributed light 
to dark brown pigmentation or may have dark 
margins with light brown or pale between; inter- 
orbital bar and principal saddle-like markings are 
distinct in 14-mm fish. 


REMARKS: Gilbert (1890) failed to select a holo- 
type and based his description on specimens 
from both Atlantic (GRAMPUS Sta. 5108, 5112) 
and Pacific waters (ALBATROSS Sta. 2827, 2829). 
This ultimately resulted in uncertainty (Bohlke, 
1968; Dawson, 1974) as to which of the two in- 
cluded species should be considered semicinctus, 
the type-species of Gillellus. | have recently 
found that no such problem exists and that the 
species must be based on Pacific material. Jordan 
(1897) referred to Gillellus semicinctus (p. 229) 
stating, “I present a figure of this interesting 
species from one of the types” and illustrated a 
specimen in Plate XXXII. The introduction (p. 20) 
indicates that Jordan’s material was in the Stan- 
ford collection and the list of plates (p. 244) 
states, ““Gillellus semicinctus: type. Gulf of Cali- 
fornia.’ Pacific specimens of the syntypic series 
include one from ALBATROSS station 2827 (US 
NM 126880) and two (CAS-SU 31) from ALBA- 
TROSS station 2829, located off Cabo San Lucas, 
Baja California Sur. Included in USNM 126880 is 
a handwritten label by Barton A. Bean (‘B. A. B.’’) 
dated January 1918 stating, ‘‘According to Dr. 


Jordan this is a cotype or paratype. The type is at 
Stanford. A third specimen (Florida) is in Nat. 
Museum.” 


There are two fish (25.8 and 27.6 mm SL) and 
a metal tag imprinted “drawn” in CAS-SU 31. The 
larger specimen has a total of 41 dorsal-fin sup- 
ports, whereas the smaller has 42. Jordan’s figure 
is very poor and shows a continuous rather than 
an anteriorly interrupted dorsal fin but dorsal-fin 
supports total 41. | conclude that Jordan figured 
the larger fish and thereby designated this speci- 
ment as the lectotype of Gillellus semicinctus in 
accord with Article 74b, International Code of 
Zoological Nomenclature. The paralectotype has 
been renumbered CAS-SU 67760. 


Considerable study material is available from 
Baja California waters, but representation from 
other areas is minimal and most specimens are 
small (under 30 mm SL) and in poor condition. 
Total material from the Revillagigedos consists 
of 17 specimens (10-22 mm SL) and there are 
but 8 (15-31 mm) from the Galapagos Islands. 
Available data (Table 11) suggest that counts of 
dorsal and anal rays, vertebrae, and lateral-line 
scales average slightly higher in the Baja Cali- 
fornia population (including Isla Guadalupe) than 
those of Revillagigedos or Nayarit, Mexico or 
Galapagos material. These differences may indi- 
cate distinct populations or merely clinal varia- 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


G. semicinctus 
G. chathamensis 
G. searcher! 


= G ornatus 


A G arenicola 
SG H. cinctus 


Fig. 8. General distribution maps of Pacific species of Gillellus and Heteristius. 


tion, but further treatment of this problem must 
await receipt of additional specimens. 

Among examined material, the smallest trans- 
formed male from Baja California waters was 28 
mm SL, the smallest ovigerous female was 33 
mm; a 26-mm transformed male and 7 ovigerous 
females (26-30 mm) occurred in a collection from 
Nayarit, Mexico. A 41-mm SL female contained 
224 near-ripe eggs (maximum diameter about 
0.7 mm) in the right gonad, together with about 


an equal number approximately 0.3 mm in dia- 
meter. 

Among Pacific dactyloscopids, the recorded 
bathymetric range of Gillellus semicinctus is 
matched only by that of G. arenicola and Dacty- 
loscopus lunaticus. Whereas the latter forms may 
occur in depths of a meter or less, | have no con- 
firmed records of Gillellus semicinctus from less 
than 5 m. Among collections with complete data, 
29 were by dredge and 15 with SCUBA; 8 were 


38 PROC. CALIF. ACAD. SClI., 4TH SER., VOL. 41, NO. 2 


Fe, 


Fig. 9. Gillellus chathamensis n. sp. (GCRL 13215, female paratype, 23 mm SL). 


Inéz to Isla Angel de la Guarda: CAS 31809 and 31810; 


in confirmed depths of 5-15 m, 22 in 17-46 m, 
GCRL 13246 and 13341; LACM 23982 through 23984, 23990 


and 9 in 50-137 m. Known from Islas Revillagig- 
edos, Isla Guadalupe (ca. 28°53’N) and Pacific 
coast of Baja California, western Gulf of Cali- 
fornia, various mainland and insular shores from 
Nayarit, Mexico to Isla Gorgona, Colombia and 
Galapagos Is. (Archipiélago de Colon), Ecuador 
(Fig. 8). 


MATERIAL EXAMINED: Two hundred eighty-two specimens, 
9.6-48.3 mm SL, including lectotype and two paralectotypes. 

Lectotype. —CAS-SU 31 (27.6 mm SL, male), Mexico, Baja 
California Sur, 22°52'00'’'N, 109°55’'00’’W, 56.7 m, 1 May 
1888, ALBATROSS Sta. 2829 

Paralectotypes. —-CAS-SU 67760 (25.8 mm), taken with 
lectotype. USNM 126880 (ca. 31.2 mm), Mexico, Baja Cali- 
fornia Sur, 24°11'45’"N, 109°55’00""W, 18.3 m, ALBATROSS 
Stan 28277, 

Other Material. —MEXICO, Baja California Norte (Pacific). 
—Isla Guadalupe: SIO 50-31, 57-190, 58-497 and GCRL 13241. 
—Isla Cedros: SIO 63-168, 71-33 and LACM 23999. Baja Cali- 


fornia Sur (Pacific). —Bahia Magdalena to Cabeza Ballena: 
LACM 23985, 23986, 23996, 23997 and SIO 64-61. Baja Cali- 
fornia Sur (Gulf of California). —Punta de los Frailes to Punta 


Concepcion: CAS 32037; CAS-SU 5934; GCRL 13242 through 
13245; LACM 9728-2, 23987 through 23989, 23994 and 23998: 
SIO 61-247, 61-249, 65-243, 65-270, 65-278, 65-295, 65-311, 
65-317, 65-321, 65-335 and 65-337; UCLA W65-49 and W65- 
86. Baja California Norte (Gulf of California). —Isla Santa 


through 23992, and 23995; SIO 65-306; UCLA W53-69 and 
W53-73. Islas Revillagigedos. —Isla Socorro: SIO 70-392. Isla 
Clarion: UCLA W55-136. Nayarit to Oaxaca: CAS 6923, 32035 
and 32036; LACM 23993. 

COSTA RICA. —Puntarenas: UCR 664-6. —Isla del Cano: 
GCRL 13247 and 13248; LACM 32547-48 and 32562-46, UCR 
675-11 and 685-1. 

PANAMA. —Islas Secas: USNM 101734. —Isla Jicaron: SIO 
71-87. 

COLOMBIA. —LACM 23981 and USNM 101725. —Isla Gor- 
gona: USNM 101738. 

ECUADOR, Galapagos Islands (Archipiélago de Colon). 
—Isla Isabela (Albemarle Is.): LACM 23980. —Isla Santa Maria 
(Charles Is.): UCLA W67-43 and USNM 101735. —Isla Baltra 
(S. Seymour Is.): USNM 109426. -Isla Santa Cruz (Inde- 
fatigable Is.): CAS-SU 39962. —Isla Marchena (Bindloe Is.): 
USNM 205457. 


Gillellus chathamensis Dawson, new species 
(Figure 9.) 


DIAGNOSIS: Dorsal finlet 3-spined, without iso- 
lated spines between finlet and remainder of 
dorsal fin; pectoral fin-rays modally 13; seg- 
mented anal rays modally 34; straight lateral-line 
scales 23-25 (usually 24); eye flap usually obso- 
lete or vestigial; without paired pores in infra- 
orbital canal. 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


DESCRIPTION: Measurements (mm) of 25.8-mm 
SL transformed male holotype follow: caudal fin 
length 3.8; length of uppermost segmented 
caudal ray 3.0; length of lowermost segmented 
caudal ray 2.7; depth of caudal peduncle 1.1; 
body depth 2.8; predorsal length 4.5; preanal 
length 6.8; pelvic to anal distance 3.0; head 
length 5.2; head breadth 2.8; maxillary to upper 
opercular angle 4.8; maxillary to upper preoper- 
cular angle 3.4; length of anterior naris 0.2; eye 
diameter 1.2; postorbital length 3.0; snout length 
1.2; maxillary length 1.8; length of 1st dorsal 
spine 1.5, 3rd spine 0.5, 4th spine 0.9, last spine 
1.5; length of 1st segmented dorsal ray 1.9. See 
Tables 2-8 for meristic data. 

Lower jaw slightly protruding, rounded to sub- 
triangular in front; 4 fimbriae on lower lip; pre- 
orbital short, moderately inclined; tubiform an- 
terior naris located near anterior rim of preorbi- 
tal; eye flap usually vestigial or obsolete; height 
of dorsal finlet about equals that of remaining 
spinous dorsal fin, somewhat less than length of 
longest segmented dorsal ray. Pectoral-fin rays 13 
in 87% of 134 counts, one fish had 12 rays in 
each fin, whereas three had bilateral counts of 
14; pectoral fins not broadly expanded in trans- 
formed males, may reach past lateral-line de- 
flection; tip of adpressed fin reaches vertical be- 
tween dorsal-fin elements 17 and 22 in trans- 
formed males, between 14 and 18 in young and 
females; segmented caudal rays frequently 
simple, but some median rays branched in most 
material examined (Table 9); opercular fimbriae 
4-8 (Fig. 7; Table 10); infraorbital canal pores 7, 
none paired. 

Arched portion of lateral line deflects between 
verticals from dorsal-fin elements 17 and 21, usu- 
ally between 18 and 20; squamation above lat- 
eral-line arch similar to that of G. semicinctus; 9 
longitudinal scale rows cross vertical near origin 
of straight lateral line; squamation apparently 
complete in 16-mm fish. Dentition essentially as 
in| semicinctus; pseudobranchiae 4-5 in two 
specimens examined; proximal pterygiophore of 
1st dorsal spine broad. 


Color in alcohol. —Ground color is pale 
in all material, markings tan to dark brown. 
Usually with only faint indication of an 
interorbital bar and without suborbital bar 
to angle of gape; 4-5 prominent spots about 
eye and with additional spots on upper sides and 
dorsum of head, some fish with a rather large 
brownish blotch on dorsum immediately behind 
each eye; usually with a faint narrow bar cross- 


1139) 


ing upper and lower jaws on either side of sym- 
physis and 2-4 isolated spots on ventral margins 
of lower jaw; there is a small blotch dorsad be- 
fore 1st dorsal spine and 7-8 (usually 7) saddle- 
like primary bars cross dorsum from finlet to 
caudal-fin base, margins of bars often incom- 
pletely pigmented and median areas poorly 
marked; secondary bars often indicated by spots 
or faint blotches on upper portions of pale inter- 
spaces; head and sides elsewhere pale. Dorsal 
finlet usually with some brown pigmentation, 
this may be restricted to posterior membrane, to 
a few isolated spots, or may shade entire finlet; 
other fins immaculate. Specimens from SEARCH- 
ER Sta. 517 are more strongly marked and ap- 
proach G. semicinctus in general coloration. 
They have prominent interorbital and suborbital 
bars, there is little or no pigment on snout or 
jaws, the predorsal spot (if any) is included in 
the 1st (anterior) primary bar, primary bars are 
more or less evenly pigmented and some fish 
have a few brown flecks anteriad above anal fin 
base. 


ETYMOLOGY: The name chathamensis refers to 
the type-locality, Chatham Bay, Isla del Coco. 


COMPARISONS: This species is distinguished 
from the closely related G. semicinctus by modal 
counts of 13 pectoral rays and 24 straight lateral- 
line scales (12 and 19-20 in semicinctus). The 
combination of unpaired infraorbital canal pores, 
3-spined dorsal finlet, and modally 13 pectoral 
rays and 24 straight lateral-line scales separates 
Gillellus chathamensis from all Pacific congeners. 


REMARKS: This species may readily be confused 
with G. semicinctus, and careful examination is 
required for identification. Among material ex- 
amined, the smallest transformed male is 18.9 
mm SL, the smallest ovigerous female (19.6 mm) 
contained 104 developing eggs, about 0.5 mm in 
maximum diameter. 


All specimens have been taken with SCUBA in 
depths of 5-12 m. Gillellus chathamensis is 
known only from Isla del Coco, Costa Rica (Fig. 
8). 


MATERIAL EXAMINED: Holotype and_ sixty-six paratypes, 
15-32 mm SL. 

Holotype. —LACM 32256-23 (25.8 mm SL, male), Costa Rica, 
Isla del Coco, Chatham Bay, 05°33'12’’N, 87°02'14’'W, 6 m, 
SCUBA, R/V SEARCHER, sta. 511, 1 April 1972, R. J. Laven- 
berg and party. 

Paratypes. —COSTA RICA, Isla dei Coco, Chatham Bay: 
UCR 711-16, GCRL 13215 and 13216, 05°33'29’’N, 87°02'48''W, 
11 m, SCUBA, R/V SEARCHER Sta. 509. LACM 32260-22, 


140 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


Fig. 10. Gillellus arenicola Gilbert (GCRL 13240, 34 mm SL, young male). 


GGRE1382135705233121NF 87-°03/157W, 12.m; SGUBA, IR/V 
SEARCHER Sta. 517. LACM 32256-24, GCRL 13211 and 13212, 
taken with holotype. —Wafer Bay: LACM 32272-18, GCRL 
13214, 05°32’45’’N, 87°03'49"W, 5 m, SCUBA R/V 
SEARCHER Sta. 532. 


Gillellus arenicola Gilbert 
(Figure 10.) 


Gillellus arenicola Gilbert, 1890: 99 (original description; 
Cape San Lucas, Baja California Sur). 
DIAGNOSIS: Dorsal finlet 2-spined; pectoral fin- 
rays modally 12; scales in straight lateral line 27- 
29 (x = 28); lateral line deflects between verti- 
cals from dorsal-fin elements 18 and 22; lower 
jaw fleshy and strongly protruding (in undamaged 
fish); without paired pores in infraorbital canal. 


DESCRIPTION: Measurements (mm) of 41.1-mm 
SL neotype follow: caudal-fin length 7.5; length 
of uppermost segmented caudal ray 5.5; length 
of lowermost segmented caudal ray 5.0; depth of 
caudal peduncle 2.0; body depth 4.6; predorsal 
length 7.7; preanal length 11.5; pelvic to anal 
distance 3.9; head length 8.9; head breadth 3.6; 
maxillary to upper opercular angle 8.0; maxillary 
to upper preopercular angle 5.8; length of an- 
terior naris 0.2; eye diameter 1.6; postorbital 
length 5.4; snout length 2.1; maxillary length 
2.8; length of 1st dorsal spine 0.9, 2nd spine 


0.7, 4th spine 0.6, last spine 2.1. See Tables 2-8 
for meristic data. 

Lower jaw distinctly protruding and produced 
anteriad as a conical fleshy prominence (Fig. 5); 
labial fimbriae 4; preorbital long, about equals 
diameter of pigmented eye, not steeply inclined; 
tubiform anterior naris located near middle of 
preorbital length, well removed from anterior 
rim; eye usually with a flap. Height of dorsal fin- 
let about 1/3 to 1/4 that of spinous portion of 
continuous dorsal fin; dorsal fin slightly notched 
or emarginate between last spines and anterior 
segmented rays. Finlet typically 2-spined (3- 
spined in two of 58 examined) and with one or 
two (40% of 35 examined) isolated spines be- 
tween finlet and continuous dorsal fin (Fig. 6). 
Pectoral fin-rays 12 in 75% of 89 counts, 7 fish 
with bilateral counts of 13; pectoral membranes 
expanded and the fin elongate in transformed 
males, may reach to or somewhat beyond origin 
of straight lateral line; tip of adpressed fin 
reaches vertical between dorsal-fin elements 15 
and 20 in transformed males, between 12 and 17 
in young and females; some median caudal rays 
branched in adults (Table 9); opercular fimbriae 
3-8 (Fig. 7; Table 10); infraorbital canal pores 
usually 7, none paired, 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


141 


TABLE 3. Frequency distributions of segmented anal and pectoral fin-rays in Pacific species of Gillellus, Sindoscopus and Heteristius. 
S Segmented anal Gane : ea 
Genus and species N 30 31 32 33 34 35 36 37 38 39 40 41 x 
Gillellus 
semicinctus 219 4 17 64 100* 31 3 332,7/ 
chathamensis 56 So PD 7ET ROSS aei3 34.5 
ornatus 75 2 Oe sbi 923 5 B5r2 
arenicola 52 1 We eas 6 1 35.8 
searcheri lO S22 25 48 Bile 
Sindoscopus 
australis 89 3 Bo se WC 28a, 
Heteristius 
cinctus 107 PANG I 7/ 2 34.6 
Pectoral fin-rays 
N 1 12 13 14 15 x 
Gillellus 
semicinctus 437 U2 3A ANS 2 Pel 
chathamensis 134 5 Wee 13 13511 
ornatus 150 4 139* 7 12.0 
arenicola 89 O° 2D 12.2 
searcheri 306 Assy |G 1301 
Sindoscopus 
australis 185 37 148* 13.8 
Heteristius 
cinctus 208 eA e2.7, 14.0 


*Primary type. 


Arched portion of lateral line deflects between 
verticals from dorsal-fin elements 14 and 17, usu- 
ally between 15 and 16; area above arch usually 
fully scaled in adults, 3-4 scale rows between 
base of 1st dorsal spine and 1st or 2nd lateral- 
line scale, narrowing to a single row above pos- 
terior half of arch (Fig. 6); 9 longitudinal scale 
rows cross vertical near origin of straight lateral 
line; lateral-line scales present in 14-mm SL fish, 
squamation apparently complete in some 22-mm 
specimens. Dentition essentially as described for 
G. semicinctus; pseudobranchiae 4 (2 examined); 
proximal pterygiophore of 1st dorsal spine com- 
paratively slender (Fig. 3). 

Color in alcohol. —Coloration is poorly pre- 
served in study materials but well marked speci- 
mens have indications of an interorbital bar and 


a more or less distinct suborbital bar crosses 
posterior third of gape to terminate on ventral 
margin of lower jaw; a narrow bar crosses under- 
side of lower jaw; upper sides and dorsum of 
head plain, lightly spotted or blotched with 
brown. Dorsum crossed by 7 principal saddle- 
like bars between nape and caudal-fin base; late 
juveniles and adults often with secondary bars 
which may approach size and pigmentation of 
primary bars, the net effect being a series of 12- 
14 saddles; pale interspaces of upper side some- 
times flecked with brown and a few scattered 
flecks may be present on lower portion of side; 
all fins immaculate. 

COMPARISONS: The 2-spined dorsal finlet and 
long, distally fleshy, lower jaw separate undam- 
aged Gillellus arenicola from all Pacific con- 


142 PROG. CALIF; ACAD. SGI., 48H SER., VOL. 47,(\NO: 2 


geners. Specimens with damaged lower jaw can 
be confused with G. ornatus, the only other spe- 
cies with 2-spined finlet (see ornatus for further 
comparisons). It is distinguished from searcheri 
by a number of meristic characters (Tables 2-8) 
and from both semicinctus and chathamensis by 
modal frequency of total dorsal spines (14 in 
arenicola against 13), number of straight lateral- 
line scales (27-29 against 18-25), and position of 
lateral-line deflection (between dorsal-fin ele- 
ments 14 and 17 versus 17 and 24 in semicintus 
and chathamensis). The several Pacific species of 
Myxodagnus (see Dawson, 1976) also have 
projecting lower jaws and are superficially similar 
to Gillellus arenicola. In Myxodagnus, however, 
there are both upper and lower labial fimbriae, 
dorsal fin is continuous and originates near verti- 
cal from anal-fin origin, and there are no promi- 
nent saddle-like blotches crossing the dorsum. 


x 
29.0 
28.2 
28.8 
25.4 
yi? 
33.0 
21.4 


62 


Ces M4 SS 7 ace 


1 


1 


1 


1 


REMARKS: Gilbert’s (1890) description was based 
on “A single specimen, 1-1/2 inches long, from 
Cape San Lucas.” This fish was not illustrated; 
there is no record of its receipt at the National 
Museum, and it is apparently lost. | therefore 
select a 41-mm SL specimen (CAS-SU 67821) as 
the neotype of Gillellus arenicola Gilbert. This 
fish, apparently an immature male, was collected 
near the type-locality; see measurements above, 
material examined and Tables 2-8 for additional 
data. 

There are relatively few specimens in collec- 
tions and most of these are in rather poor con- 
dition. Among material examined, the smallest 
transformed male is 32 mm SL, the smallest ovig- 
erous female 40 mm; a 49-mm female contained 
129 developing eggs in the right gonad. Gillellus 
arenicola has been taken with seine, dredge or 
SCUBA to depths of 137 m. It is known from the 
southern portion of Baja California Sur and from 
single collections off the states of Colima and 
Oaxaca, Mexico (Fig. 8). 


sy eisk eb sky si yf sis) Ss tas} yh bi sh te) YS). 


Number of arched lateral-line scales 
1 


19 BO Bi) D2 PB BA Wh A Py/ Ph} VE) 30) Sil 
1 
ies (7 Ia) 24 Gt 


810 9 4 9 3 


M7 39) bye 2 
3\* 
A137" 22 


5 §) 70) 20 7/ 2 © 


1 
1 
1 


Frequency distribution of arched lateral-liné scales in Pacific species of Gillellus, Sindoscopus and Heteristius. 
3} iI) 17/7 


18 
1 


MATERIAL EXAMINED: Fifty-eight specimens, 13-50 mm SL, 
including neotype. 


Neotype. —CAS-SU 67821 (41.1 mm SL), Mexico, Baja 
California Sur, Bahia de San Lucas, 22°53’00’’N, 109°52’00’’W, 
Sefton-Stanford ORCA Expdn. Sta. 44-D-2, 19 May 1952. 


Other Material. —MEXICO, Baja California Sur. —San Jaime 
Bank to Bahia de las Palmas: AMNH 5398 and BC 60-513; 
CAS 2716, 14212 and 32038 through 32040; CAS-SU 18915; 
GCRL 13237 through 13240; LACM 23968, 23969 and 31774- 
45; SIO 61-232, 61-247 and 62-704 or 59-210 (loc. uncer- 
tain); UCLA W59-251; and USNM 2533. Colima: LACM 
23967. Oaxaca: CAS 32034. 


N 
87 
25 
47 
29 
93 
62 
49 


TABLE 4. 


cinctus 


semicinctus 
chathamensis 
ornatus 
arenicola 
searcheri 
australis 
*Primary type. 


Genus and species 
Gillellus 
Sindoscopus 
Heteristius 


143 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


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SNYPPT!D 
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144 


PROG. GALE AGAD® SGI 4h SERY VOL 41 NOF2 


Fig. 11. Gillellus ornatus Gilbert (GCRL 13234, 41 mm SL, transformed male). 


Gillellus ornatus Gilbert 
(Figure 11.) 


Gillellus ornatus Gilbert, 1892: 558 (original description, 
Gulf of California). 


DIAGNOSIS: Dorsal finlet 2-spined; pectoral fin- 
rays modally 12; scales in straight lateral line 22- 
26 ( mean = 24); lateral line deflects between 
verticals from dorsal-fin elements 18 and 22; 
lower jaw neither fleshy nor strongly protruding 
anteriad; without paired pores in_ infraorbital 
canal. 


DESCRIPTION: Measurements (mm) of 36.2-mm 
SL transformed male neotype follow: caudal fin 
length 6.4; length of uppermost segmented 
caudal ray 5.0; length of lowermost segmented 
caudal ray 4.3; depth of caudal peduncle 1.7; 
body depth 3.6; predorsal length 6.2; preanal 
length 9.2; pelvic-anal distance 3.4; head length 
7.5; head breadth 3.5; maxillary to upper oper- 
cular angle 7.1; maxillary to upper preopercular 
angle 4.7; length of anterior naris 0.2; eye dia- 


meter 1.5; postorbital length 5.1; snout length 
0.6; maxillary length 2.0; length of 1st dorsal 
spine 1.3, 2nd spine 0.8, 3rd spine 0.6, 4th spine 
1.7, last spine 1.8; length of 1st segmented dor- 
sal ray 2.2. See Tables 2-8 for meristic data. 


Lower jaw slightly protruding, not continued 
as a fleshy prominence anteriad; labial fimbriae 
4; preorbital short, moderately inclined; tubi- 
form anterior naris slightly nearer anterior rim 
than middle of preorbital length; eye usually 
with a prominent flap. Height of dorsal finlet 
usually less than that of continuous portion of 
dorsal fin, segmented dorsal rays longer than 
spines. Finlet always 2-spined, with one (95%) 
or two isolated spines between finlet and con- 
tinuous portion of dorsal fin (Fig. 6); membrane 
rather deeply incised between 1st and 2nd spines 
of continuous dorsal fin, the fin emarginate be- 
tween last spine and 1st segmented ray. Pectoral 
fin-rays 12 in 93% of 150 counts, one fish with 
11 rays in each fin, three with bilateral counts of 
13; pectoral fin elongate in transformed males, 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


TABLE 7. Frequency distributions of caudal vertebrae and opercular fimbriae in Pacific species of Gillellus, 


145 


Sindoscopus and 


Heteristius. 
Caudal vertebrae i 
Genus and species N 30 31 32 33 34 35 36 37 38 39 40 41 42 x 
Gillellus 
semicinctus 190 Se MO 57 85s 24 88} 05 
chathamensis 56 3 RH 3} 3 B55 
ornatus 72 1 23 eS Ona 35.9) 
arenicola 47 O28 eo 1 36.0 
searcheri 198 Yq Bil [ses SX) 2 32.0 
Sindoscopus 
australis 93 26 43 Di | 3 40.0 
Heteristius 
cinctus 134 (HO G0) 7 3535 
Opercular fimbriae 
N 2 3 4 5 6 7 8 9 10 Th 12 
Gillellus 
semicinctus 365 4 i 58* 109* 102 55 27 2 1 
chathamensis 11333) 3 BS) ZIG 
ornatus 146 1 1 4 ai sty 32 13) 4 if 
arenicola 79 1 Ye DRY SNF] 
searcheri 318 1 SS OZ 658 8 
Sindoscopus 
australis 180 2 1) LO 5.6 28 4 1 
Heteristius 
cinctus DiV7 1 1 Sy (7/3 GF 1 
*Primary type. 
the interradial membranes expanded, the fin scale, narrowing to a single crowded row behind 


somewhat cupped or outwardly convex; tip of 
adpressed fin falls short of origin of straight 
lateral line, reaches vertical between dorsal-fin 
elements 15 and 19 in transformed males, be- 
tween 12 and 15 in young and females; caudal fin 
formula modally 2 + 6 + 2 in examined mate- 
rial (Table 9); opercular fimbriae 2-10 (Fig. 7; 
Table 10); infraorbital canal pores 6-7, none 
Paired. 

Arched portion of lateral line deflects between 
verticals from dorsal-fin elements 18 and 22, 
usually between 18 and 21, squamation variable 
above lateral-line arch; naked in early juveniles 
and some adults; some with a few scattered 
scales anteriad (Fig. 6), others with about two 
rows originating above 2nd or 3rd lateral-line 


and replaced by impinging lateral-line scales over 
posterior half of arch; 7-9 longitudinal scale 
rows cross vertical near origin of straight lateral 
line; arch scales present in 12-mm SL fish, squa- 
mation apparently complete in some 19-mm 
specimens. Jaw teeth apparently in two rows 
over anterior half of upper jaw and in 2-3 rows 
near symphysis of lower jaw, teeth in single row 
posteriad; pseudobranchiae 3-4 (3 examined); 
proximal pterygiophore of 1st dorsal spine broad 
(Fig. 3). 

Color in alcohol. —An interorbital bar present 
and continued across suborbital as a broad 
blotch which may extend around and below 
posterior angle of gape to posteroventral margin 
of lower jaw; a faint narrow bar extends across 


146 


upper and lower jaw on either side of symphysis; 
upper sides and dorsum of head blotched and 
spotted with brown, head elsewhere mainly pale. 
Dorsum crossed by 8-10 (usually 9) principal 
saddle-like bars between nape and caudal-fin 
base; well marked late juveniles and adults have 
secondary bars, most distinct near dorsal-fin 
base, but frequently indicated only by brown 
flecks on upper sides; ventral margins of primary 
bars (behind nape) may be more or less united 
by irregular brown flecks along midline of sides; 
body elsewhere mainly pale, but some specimens 
lightly flecked with brown on lower sides; fins 
mainly pale but occasionally there is a trace of 
brown shading on dorsal finlet; primary bars are 
distinct in 12-mm fish. 


COMPARISONS: Among Pacific congeners, Gill- 
ellus ornatus shares the 2-spined dorsal finlet 
only with the sympatric G. arenicola. It differs 
from that species in average counts of arched 
and straight lateral-line scales (29 and 24 versus 
25 and 28 in arenicola) and modal counts of 
total dorsal spines (13 against 14). In addition, 
the lateral-line arch deflects between verticals 
from dorsal-fin elements 18 and 22 in ornatus 
(between 14 and 17 in arenicola) and lower jaw 
is short and scarcely protruding (distally fleshy 
and protruding in undamaged arenicola). A num- 
ber of meristic characters (Tables 2-7) distinguish 
ornatus from the aberrant G. searcheri but the 
remaining 3-spined species (semicinctus and 
chathamensis) may be confused with ornatus 
wherein the finlet has been damaged. In such 
cases, Gillellus ornatus is best separated from 
chathamensis by the lower modal count of 
pectoral rays (12 against 13 in chathamensis), 
and from semicinctus by higher average values 
for total dorsal-fin elements and straight lateral- 
line scales (44 and 24 versus 41 and 20 in semi- 
cinctus). 


REMARKS: Gilbert (1892) based his description 
on a single specimen (“about 2 inches long’) 
from ALBATROSS Sta. 2828 located off the W 
side of Isla Cerralvo in the Gulf of California 
(24°11'30’"N, 109°55’00’W). This specimen was 
never received at the National Museum, has 
never been illustrated and is presumably lost. 
| therefore select a mature male (LACM W65-86- 
1) as the neotype of Gillellus ornatus Gilbert 
(see measurements above, material examined 
and Tables 2-8 for additional data). 

Among material examined, the smallest trans- 
formed male is 28.0 mm SL, the smallest oviger- 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


ous female 30.6 mm; a 51-mm female contained 
214 near-ripe eggs, maximum diameter about 
0.9 mm. 

Available data suggest that all collections have 
been made with dredge or SCUBA over a 3-55 m 
depth range. Gillellus ornatus is known only 
from the Gulf of California between Los Frailes 
and Isla Angel de la Guarda in the west and 
from a single eastern Gulf collection near Guay- 
mas, Sonora (Fig. 8). 


MATERIAL EXAMINED: Ninety-seven specimens, 12-51 mm 
SL, including neotype. 

Neotype. —LACM W65-86-1 (37.0 mm SL, male), Mexico, 
Baja California Sur, Isla Santa Cruz, 22 July 1965, B. W. 
Walker and party. 

Other Material. —MEXICO, Gulf of California. —Baja Cali- 
fornia Sur: CAS 24624 and GCRL 13234 through 13236; 
LACM 23964 and 23966; SIO 61-247 through 61-249, 65-295, 
65-311 and 65-323; UCLA W65-71 and W65-86. —Baja Cali- 
fornia Norte: LACM 23965, UCLA W53-73 and W53-86. 
—Sonora: SIO 70-84. 


Gillellus searcheri Dawson, new species 
(Figure 12.) 


DIAGNOSIS: Dorsal finlet 3-spined, without iso- 
lated spines between finlet and remainder of 
dorsal fin; dorsal spines total 20-23; pectoral fin 
rays modally 13; scales in straight lateral line 9- 
13; lateral line deflects between verticals from 
dorsal-fin elements 27 and 32; eye flap present; 
paired pores in posterior portion of infraorbital 
canal. 


DESCRIPTION: Measurements (mm) of 24.5-mm 
SL transformed male holotype follow: caudal fin 
length 4.4; length of uppermost segmented ray 
3.8; length of lowermost segmented ray 3.6; 
depth of caudal peduncle 1.5; body depth 2.9 
predorsal length 4.3; preanal length 6.7; pelvic to 
anal distance 2.7; head length 5.3; head breadth 
3.2; maxillary to upper opercular angle 4.8; max- 
illary to upper preopercular angle 3.4; length of 
anterior naris 0.3; eye diameter 1.1; postorbital 
length 3.1; snout length 1.0; maxillary length 
1.7; length of 1st dorsal spine 1.6, 3rd spine 0.6, 
4th spine 1.1, last spine 0.5; length of 1st seg- 
mented dorsal ray 1.4. See Tables 2-8 for meristic 
data. 

Lower jaw but slightly protruding; lower lip 
with 4-7 papilla-like fimbriae, typically 6 in late 
juveniles and adults (Fig. 5); preorbital very 
short, much less than eye diameter, steeply in- 
clined; tubiform anterior naris located close to 
anterior rim of preorbital and with a distal valve- 
like emargination; eye with a distinctive flap lo- 
lated on posterior third of eye, narrow at base 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


147 


Fig. 12. Gillellus searcheri n. sp. (GCRL 13223, male paratype, 25 mm SL). 


but distally expanded. Height of dorsal finlet 
equal to or somewhat less than maximum height 
of remainder of spinous dorsal; last dorsal spines 
of continuous dorsal fin much shorter than the 
following segmented rays, the fin deeply notched 
or emarginate between (Fig. 13). Pectoral fin- 
rays 13 in 93% of 306 counts, three fish had bi- 
lateral counts of 14; pectoral membranes some- 
what expanded and the fin elongate in  trans- 
formed males but always falls well short of origin 
of straight lateral line; tip of adpressed_ fin 
reaches vertical between dorsal-fin elements 14 
and 18 (usually between 15 and 18) in trans- 
formed males, between 12 and 17 (usually 13 
and 16) in young and females; caudal-fin formula 
modally 1 + 8 + 1 in adults (Table 9); opercular 
fimbriae 4-9 (Fig. 7; Table 10). Infraorbital canal 
typically with paired pores in lachrymal branch 
and in three posterior positions (Fig. 1), pores 
usually total 11 or 12. 

Archer portion of lateral line deflects between 
verticals from dorsal-fin elements 27 and 32, usu- 


ally between 28 and 30; area above arch mostly 
naked, adults often with 1-6 embedded scales 
anteriad on either side of dorsal-fin base (Fig. 
13); 5-7 (usually 5) longitudinal scale rows cross 
vertical near origin of straight lateral line; scales 
present in lateral-line arch of 11-mm fish, squa- 
mation apparently complete in some 15-mm-SL 
specimens. Upper jaw teeth in about three rows 
near symphysis, narrowing to a single row pos- 
teriad; lower jaw teeth mainly uniserial with but 
1-2 paired teeth on either side of symphysis; 
pseudobranchiae 4-5 (3 examined); proximal 
pterygiophore of 1st dorsal spine broad (Fig. 3). 

Color in alcohol. —Head usually without a dark 
interorbital bar but with a broad  suborbital 
blotch which may include posterior portion of 
gape and continue on ventral margin of lower 
jaw; preorbital spotted or streaked with brown; 
usually with a narrow bar crossing upper lip and 
tip of lower jaw on either side of symphysis; up- 
per sides and dorsum of head spotted, blotched 
or mottled with brown. Early juveniles with about 


148 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


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ERD DELS ID 
333 3339939232239 3999 339- 
39°? 


53279? 


passceeee ee 
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(Qo ERPs 
U8 


Fig. 13. Lateral and dorsal views illustrating lateral-line configuration and squamation 
above lateral-line arch in Gillellus searcheri (top), Sindoscopus australis (middle) and Heterist- 


ius cinctus (bottom). 


6 primary saddle-like bars crossing dorsum from 
nape to caudal-fin base and secondary bars may 
be present in 15-mm-SL specimens. In late juven- 
iles and adults, primary and secondary bars are 
largely indistinguishable in lateral aspect (Fig. 12) 
and interspaces are often heavily spotted or 
flecked with brown. In dorsal aspect, 6 primary 
and 4-5 secondary bars are indicated for most 
specimens by darkly pigmented blotches along 
dorsal-fin base; secondary bars are represented 
by a single blotch, whereas primary bars are 
identified by two blotches separated by a short 
pale interspace; dark blotches along base of con- 
tinuous dorsal fin usually total 13-14; lower sides 
pale or with a narrow stripe of brown flecks just 
above anal-fin base. Dorsal finlet with 2-3 brown 
spots on 1st spine, elsewhere usually spotted or 
shaded with brown; continuous dorsal fin with 
3-5 well spaced brown spots on most fin-rays, 
the membranes mainly pale; caudal fin-rays 


crossed by about 4 irregular vertical rows of 
spots; upper 5-6 pectoral rays crossed by a 
brown bar near their base and another near 
middle of pectoral length; pelvic and anal fins 
immaculate. Degree of pigmentation is variable 
but the individual character of the saddle-like 
bars is masked by flecks and spots in most adults 
and upper sides appear generally brownish to 
the naked eye. 


ETYMOLOGY: The name searcheri refers to the 
R/V SEARCHER. Collections from this vessel, 
owned by the Janss Foundation and destroyed by 
fire in May 1972, have contributed much to our 
knowledge of Pacific dactyloscopids. 


COMPARISONS: Gillellus searcheri differs from 
Pacific congeners in counts of total dorsal spines 
(20-23 against 11-15 in other species), segmented 
dorsal rays (16-19 against 25-33), arched lateral- 


149 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


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line scales (52-62 against 23-33) and _ straight 
lateral-line scales (9-13 against 18-29). Other spe- 
cies typically have 4 labial fimbriae in adults, an- 
terior naris is a simple tube, the eye flap (when 
present) is located near middle of eye diameter 
and there are no paired pores in the infraorbital 
canal. There are usually 6 labial fimbriae in adult 
searcheri, the anterior naris has a distal valve- 
like emargination, the eye flap is located over 
posterior third of eye and there are paired pores 
in the infraorbital canal. This species appears 
most closely related to the Atlantic G. greyae 
Kanazawa which shares the emarginate naris and 
eye-flap position, has more than 4 labial fimbriae 
and also has paired infraorbital canal pores. 
There are, however, fewer scales in the lateral- 
line arch of greyae (about 28-34) and total lat- 
eral-line scales are more numerous in searcheri 
(64-73 versus about 53-58 in greyae). 


REMARKS: Although Gillellus searcheri (and G. 
greyae) is obviously distinct from Pacific con- 
geners in several features, close relationship is 
indicated by a number of common characters 
such as general dorsal-fin configuration, number 
of principal caudal rays, the naked venter, re- 
striction of labial fimbriae to lower lip, etc. | find 
no useful purpose in erecting a separate genus 
or subgenus to accommodate this somewhat ab- 
errant species (or species group). The paired in- 
fraorbital pores suggest that searcheri is inter- 
mediate between the more typical species of 
Gillellus and Sindoscopus (described below). 


Specimens from Mexican waters are somewhat 
more heavily pigmented than Costa Rican and 
Panamanian material, and there appear to be 
minor meristic differences in these populations 
(Table 11). Among material examined, the small- 
est transformed male is 21.6 mm SL, the smallest 
ovigerous female 21.5 mm; a 24-mm female con- 
tained 41 developing eggs in the right gonad. 


Recorded to a confirmed maximum depth of 
15 m, Gillellus searcheri is known from the Islas 
Tres Marias and Nayarit, Mexico and from insular 
localities in Costa Rica and Panama (Fig. 8). 


MATERIAL EXAMINED: Two hundred fifty-six specimens, 11- 
31 mm SL, including holotype and one hundred thirty-four 
paratypes. 

Holotype. —LACM 32546-45 (24.5 mm SL, male), Costa 
Rica, Isla del Cano, 08°43’15’’N, 83°53'07’’W, 9 m, SCUBA, 
R/V SEARCHER Sta. 471, 14 March 1972, W. A. Bussing and 
party. 

Paratypes. —COSTA RICA, Isla del Cano: UCR 676-9, 
08°43'16’’"N, 83°52'50’’W, 14 m, SCUBA, R/V SEARCHER 
Sta. 474. LACM 32547-49 and GCRL 13217, 08°43'15’'N, 


PROG] GALIEVAGADSSGI>) 4 SER) VOL AlNOe2 


83°53'07''W, 15 m, SCUBA, R/V SEARCHER Sta. 472. LACM 
32546-46, GCRL 13222 and 13223, taken with holotype. 


Other Material. —MEXICO, Islas Tres Marias: GCRL 13219 
through 13221; SIO 62-25, 62-56 and 62-58. Nayarit: GCRL 
13218, SIO 62-49 and UCLA W58-5. 


PANAMA, Islas Secas: SIO 70-139. —Isla Saboga: SIO 67- 
40. 


Sindoscopus Dawson new genus 
TYPE-SPECIES: Gillellus australis Fowler and Bean, 1923. 


DIAGNOSIS: Dorsal-fin origin on nape, the fin 
continuous but anterior membranes often deeply 
incised; eye not stalked and without minute 
flaps or papillae; pectoral fin-rays modally 14; 
principal segmented caudai rays 10, some 
branched in late juveniles and adults; well de- 
veloped fimbriae on both upper and lower lips; 
head, pectoral-fin base and venter naked; prin- 
cipal preopercular canals 4, unbranched; some 
paired pores in infraorbital canal; without pre- 
dorsal bones (interneurals). 


DESCRIPTION: The genus is monotypic; see de- 
scription of australis below. 


ETYMOLOGY: sindo plus scopus; in recognition 
of my Chilean field associate, Sr. Gumersindo 
Revuelta, who collected some of the present 
specimens and was instrumental in making all of 
the recent collections available for study; scopus 
- watcher, in reference to stargazers. 


COMPARISONS: The continuous dorsal fin is 
shared with Dactylagnus, Myxodagnus and some 
species of Platygillellus (Table 1). Dorsal-fin ori- 
gin is near vertical from anal-fin origin in Dacty- 
lagnus and Myxodagnus (on nape in Sindoscopus) 
and Platygillellus has 11 rather than 10 principal 
caudal rays. Closer relationship to Gillellus seems 
indicated by similarities in body configuration, 
caudal fin morphology, orientation and extent of 
straight lateral-line scales, and paired infraorbital 
pores (searcheri group only). Sindoscopus is dis- 
tinguished by the continuous rather than inter- 
rupted dorsal fin, by development of both upper 
and lower lip fimbriae (on lower lip only in 
Gillellus) and by the presence of 4 rather than 3 
primary preopercular canals. 


Sindoscopus australis (Fowler and Bean) 
(Figure 14.) 


Gillellus australis Fowler and Bean, 1923: 23 (original de- 
scription; Valparaiso, Chile). 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 151 


Fig. 14. Sindoscopus australis (Fowler and Bean). Top: GCRL 13399 (64 mm SL, male) 
from Algarrobo, Chile. Middle and bottom: GCRL 13404 (71 mm SL, transformed male) from 
Antofagasta, Chile. 


52 
AeA re ee H. cinctus 
S. australis 
W 
< 
fea 
a 
2 
ve 
a 
a 
or 
WJ 
a 
o 
= 


0 20 40 60 80 100 
STANDARD LENGTH (MM) 


Fikes 5) 


DIAGNOSIS: Diagnostic characters are those of 
the genus. 


DESCRIPTION: Measurements (mm) of 75.8-mm- 
SL male (GCRL 13399) follow: caudal fin length 
10.7; length of uppermost segmented ray 10.3; 
length of lowermost segmented ray 9.4; depth of 
caudal peduncle 3.6; body depth 8.3; predorsal 
length 13.2; preanal length 19.9; pelvic to anal 
distance 9.0; head length 14.0; head breadth 6.8; 
maxillary to upper opercular angle 12.8; maxil- 
lary to upper preopercular angle 8.9; eye dia- 
meter 2.0; postorbital length 8.7; snout length 
1.8; maxillary length 3.9; length of 1st dorsal 
spine 1.6, 2nd spine 1.6, 3rd spine 1.5, 4th spine 
1.6, last dorsal spine 2.3; length of 1st segmented 
dorsal ray 2.7. See Tables 2-8 for meristic data. 
Body rather slender, broader and deeper in 
front, more or less oval in cross section and 
gradually narrowing caudad; head flattened to 
slightly rounded dorsad, broadest near posterior 
margin of preopercle, snout not strongly in- 
clined; head, in dorsal aspect, somewhat tri- 
angular anteriad, distinctly narrowed in front 
(Fig. 5); lower jaw moderately protruding, 
rounded in dorsal profile, not continued anteriad 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


40 


30 


LOWER LIP FIMBRIAE 
DY) 
e) 


O 20 40 60 80 100 
STANDARD LENGTH (MM) 


Regressions of frequencies of labial fimbriae in Heteristius cinctus and Sindo- 
copus australis. See Table 10 for supporting data. 


as a conical, fleshy projection; upper and lower 
lips with well developed simple fleshy fimbriae, 
most numerous below, frequencies ontogenetic 
(Fig. 15; Table 10); preorbital length about 2/3 
eye diameter; eye dorsolateral, protruding but 
not on long protrusile stalk, without minute 
flaps or papillae; anterior naris a very short tube 
with distal valve-like emargination, located about 
midway between eye and anterior rim of pre- 
orbital; posterior naris a simple pore with slightly 
raised rim, located behind anterior naris and ad- 
jacent to anterior rim of orbit; principal preoper- 
cular canals 4 (Fig. 1), not branched, the 1st (an- 
terior) slightly shorter than remainder and usually 
with a single pore distad, occasionally with 1-3 
smaller pores between this and base of canal; 
infraorbital canal typically with single pores an- 
teriad, single or paired pores in  lachrymal 
branch, and paired (occasionally 3) pores in 
three posterior positions; opercle with a short 
fringe of up to 10 unbranched fimbriae overlying 
upper posterior margin, frequency ontogenetic 
(Fig. 7; Table 10); opercles broadly expanded, 
the membranous margins cover pectoral- and 
pelvic-fin bases and overlap on underside of 
head, 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


Dorsal-fin origin on nape, in advance of upper 
opercular angle (Fig. 13); dorsal fin continuous, 
anterior membranes often deeply incised; an- 
terior 4-5 spines short and rather flexible, 1st 
and 2nd more closely spaced than 2nd and 3rd; 
last dorsal spine somewhat shorter than 1st seg- 
mented ray, the fin somewhat notched or emar- 
ginate between; anal spines 2, about equal in 
length and a third shorter than 1st segmented 
ray; anterior anal-fin elements modified in trans- 
formed males, spines angled caudad, 1st seg- 
mented ray angled forward (about 45°), the 
2nd ray more nearly vertical, modified elements 
somewhat swollen or fleshy and membranes are 
stretched or elongate between. Posteriormost 
dorsal and anal rays usually free from peduncle, 
occasionally bound by short membranes; last 
anal ray reaches past rear of hypural, last dorsal 
ray somewhat shorter; caudal peduncle entire, 
without notch in ventral margin (Fig. 2); caudal 
fin rounded, upper and lowermost rays typically 
simple, subequal in length or lowermost the 
shorter; 6-9 median rays branched in study mate- 
rial, modal formula 1 + 8 + 1 (83% of 84 speci- 
mens examined). 


Pectoral fin-rays 14 in 80% of 185 counts, 11 
fish with bilateral counts of 13; pectoral fin 
rounded to somewhat pointed, 5th-7th rays from 
dorsalmost the longest, rays gradually shorten 
ventrad but not appreciably swollen or thick- 
ened; pectoral fin somewhat elongate in trans- 
formed males but usually falls short of origin of 
straight lateral line, the interradial membranes 
broad and fin cupped or outwardly convex; tip 
of straightened and adpressed fin reaches vertical 
between dorsal-fin elements 15 and 20 (usually 
16 and 19) in transformed males, between 14 
and 17 in young and females; pelvic fin I, 3; tips 
of segmented rays free with emarginate mem- 
branes between, longest ray seldom reaches anal- 
fin origin; all fin rays simple, except for branched 
caudal rays in late juveniles and adults. 


Lateral line continuous, originates above and 
in advance of upper opercular angle (on or in 
advance of vertical from 1st dorsal spine), arched 
upward to follow near dorsal-fin base, deflects 
between verticals from dorsal-fin elements 14 and 
18, and continues midlaterally to terminate as 
penultimate scale on caudal-fin base; canal of 
straight lateral-line scales parallels longitudinal 
body axis, last scale not angled ventrad (Fig. 2). 
Scales thin, more or less rounded and persistent 
in upper portion of lateral-line arch; scales of 
descending arch and straight lateral line some- 


153 


what deciduous; canal of straight lateral-line 
scales without terminal branch anteriad, often 
with a short dorsal branch and terminal pore 
on anterior third of canal. Squamation variable 
above lateral-line arch, more or less fully scaled 
or with patches of embedded scales and naked 
areas between (Fig. 13); usually with small em- 
bedded scales beginning above 2nd or 3rd later- 
al-line scale about three rows between arch and 
base of 4th or 5th dorsal spine and with two 
crowded rows of larger scales above posterior 
half of elevated arch; head, pectoral-fin base, 
venter and sides (below line from upper pectoral 
angle to near 1st segmented anal ray) naked; 11- 
13 longitudinal scale rows cross vertical near 
origin of straight lateral line. 


Premaxillaries somewhat protractile, the pedi- 
cels barely reach past middle of orbit; jaw teeth 
small, conical, somewhat curved inward, none 
distinctly enlarged; lower jaw teeth in two rows 
near symphysis, uniserial behind; upper teeth 
in three rows anteriad, two rows near middle of 
jaw and in single row behind; vomer and pala- 
tines edentate; tongue subtriangular, narrowly 
rounded in front; basihyal slender, barely spatu- 
late at tip. Three complete gill arches plus a 
hemibranch, a pore in membrane behind lower 
portion of hemibranch; no gill rakers on outer 
margin of 1st arch, 8-10 short rakers along inner 
margin; pseudobranchiae 4-6 (4 examined); 
branchiostegals 6; without predorsal bones; 
proximal pterygiophore of 1st dorsal spine not 
exceptionally broad, inserted slightly behind 1st 
neural spine (Fig. 3); abdominal vertebrae 10-11 
(usually 11) in radiographs; caudal fin with 5 
spiniform elements above and 3 below, upper 
and lower hypurals each bear 5 segmented rays 
(in 2 cleared specimens). 


Color in alcohol. —Ground color light tan to 
pale, markings tan to dark brown. Well marked 
fish with a dark interorbital bar divided below 
eye to form a short bar across posterior third of 
gape and a long narrow bar extending postero- 
ventrad to slightly beyond middle of preopercle; 
underside of lower jaw with a dark-edged blotch 
on either side of symphysis; upper sides and 
dorsum of head variously spotted or faintly 
blotched, remainder of head pale. Dorsum 
crossed by 9-10 (usually 10) dark-edged blotches 
or bars between nape and caudal-fin base; the 
anterior bar, variously solid or interrupted so as 
to form two bars with a pale area between (Fig. 
14), extends ventrad to upper margin of opercle; 
remaining bars usually fall well short of lateral 


154 


midline, seldom extending more than 3-4 scale 
rows below dorsal-fin base; usually without dis- 
tinct secondary bars, pale interspaces and area 
between ventral margins of bars and midline ir- 
regularly streaked or flecked with brown; lower 
part of side pale in some fish, others with about 
10 small blotches more or less equally spaced 
along Ist scale row above anal-fin base. Dorsal 
fin with pale membranes and 1-3 small spots on 
most fin rays; anal fin mainly pale, occasionally 
with a few isolated spots or brown shading on 
some rays and posterior membranes; pelvic fins 
pale; dorsalmost pectoral rays sometimes faintly 
spotted, the fin otherwise pale; caudal fin pale 
near base but crossed distad by about 10 vertical 
rows of small spots. 


REMARKS: Fowler and Bean (1923) described 
australis from two specimens collected by the 
Wilkes Exploring Expedition at Valparaiso, Chile. 
There have apparently been no subsequent rec- 
ords of this species and Myers and Wade (1946) 
suspected that there may have been an error in 
type locality. Present material, some of which 
was collected within 40 km of Valparaiso, clari- 
fies the matter and firmly establishes australis as 
the southernmost dactyloscopid. The holotype 
and paratype are now damaged, in very poor 
condition, and it is not possible to obtain mean- 
ingful measurements. The holotype was origin- 
ally described as having 26 dorsal spines and 21 
segmented rays, 38 segmented anal rays and 
15 pectoral rays; the lips were described as “en- 
tire.” | counted 21 spines and 25 segmented dor- 
sal rays, 39 segmented anal rays, and 14 rays in 
each pectoral fin. In addition, there are about 11 
fimbriae on the upper lip and at least 6 on the 
undamaged right side of the lower lip. Fowler and 
Bean (op. cit.) described the 6 anterior dorsal 
spines as separate but this is understandable as 
the anterior membranes are often broken (as 
they are now in the holotype) and, even in un- 
damaged specimens, membranes are low and 
may be partly concealed in the somewhat de-’ 
pressed area along anterior portion of dorsal-fin 
base. 

The bulk of recent collections have come from 
the vicinity of Valparaiso and from Antofagasta, 
some 1100 km to the north. The southern mate- 
rial averages somewhat higher in a number of 
meristic characters (Table 11), but | find no other 
substantial differences and consider such varia-, 
tion to be clinal. The upper, lower or both of the 
posteriormost spiniform caudal elements were 
segmented in all Algarrobo specimens with the 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


modal caudal formula of 1 + 8 + 1, and in 31% 
of the Antofagasta material. Segmentation of 
these typically simple elements also occurs infre- 
quently in Gille/lus but it has not been noted in 
other Pacific dactyloscopids. 

Among material examined, the smallest trans- 
formed male is 64 mm SL, the smallest ovigerous 
female 50 mm; a 74-mm female contained 419 
well developed eggs in the left gonad, maximum 
diameter about 1.1 mm. An isopod, abundant in 
the stomachs of most Algarrobo specimens, has 
been identified as /socladus sp. by Dr. Peter 
Glynn, Smithsonian Tropical Research Institute. 


DISTRIBUTION: Most material has been seined 
in depths of 2 m or less; surface temperature 
range for 3 collections was 13-14°C. Sindoscopus 
australis is known only from coastal waters of 
Chile between about 23°34’S and 33°02’S. 


MATERIAL EXAMINED: Eighty-six specimens, 32-92 mm SE 
including holotype and paratype. 

Holotype. —USNM 83315 (ca. 76 mm SL), Chile, Valpar- 
aiso, Wilkes Expdn., no other data. 

Paratype. —USNM 83099 (60.7 mm SL), data as for holo- 
type. 

Other Material. —CHILE, Antofagasta: ANSP 131422, GCRL 
13397 and 13400 through 13404, MNHN uncat. and USNM 
214226. —Coquimbo: USNM 176588. -Valparaiso: GCRL 
13398, 13399 and MNHN uncat. -Loc. uncertain; MNHN 
uncat. 


Genus Heteristius Myers and Wade 


Heteristius Myers and Wade, 1946: 160 (type-species by orig- 
inal designation, Heteristius jalisconis Myers and Wade, 
1946 [ = Dactyloscopus cinctus Osburn and Nichols]). 


DIAGNOSIS: With 2-spined dorsal finlet origi- 
nating On nape and with 1 or 2 isolated spines 
between finlet and remainder of dorsal fin; eyes 
not stalked, without flaps or papillae; pectoral 
fin-rays modally 14; principal segmented caudal 
rays 12, unbranched; fimbriae present on both 
upper and lower lips; venter scaled in late juven- 
iles and adults, head and pectoral-fin base naked; 
principal preopercular canals more than 6, un- 
branched; some paired pores in_ infraorbital 
canal; without predorsal bones (interneurals). 


DESCRIPTION: The genus is monotypic, see de- 
scription of cinctus below. 


COMPARISONS: Heteristius is a specialized 
genus, differing from other sand stargazers in 
caudal-fin morphology and in configuration of 
the preopercular canal system (Table 1). Principal 
segmented caudal rays are 12 and unbranched 
in Heteristius, whereas there are 10 or 11 in 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


Fig. 16. Heteristius cinctus (Osburn and Nichols). SIO 62-720 (31 mm SL). 


other genera and (except for a specialized spe- 
cies-group within Dactyloscopus) there is onto- 
genetic branching of the median rays. There are 
either 3 or 4 simple or distally branched princi- 
pal preopercular canals in other genera, but these 
are replaced in Heteristius by about 7-9 simple 
canals on posteroventral portion of preopercle 
(Fig. 1). The scaled venter is shared only by 
Platygillellus, but scales are here confined to a 
narrow transverse band (venter completely scaled 
in Heteristius) and the pectoral-fin base is scaled 
in adults (naked in Heteristius). 


REMARKS: Osburn and Nichols (1916) described 
Dactyloscopus cinctus from ALBATROSS collec- 
tions at Baja California Sur. Their description and 
accompanying figure indicate a continuous dor- 
sal fin and there are other errors, but the venter 
is clearly scaled and characteristic bars are shown 
on the side of the illustrated specimen. Myers 
and Wade (1946) described jalisconis (type-spe- 
cies of Heteristius) from VELERO collections off 
Jalisco and Oaxaca, Mexico. Although these au- 
thors briefly reviewed described dactyloscopids 
and made generic comparisons in their descrip- 
tion of Heteristius, they apparently overlooked 
Osburn and Nichols’ publication. | have exam- 


ined the type-material of these nominal species 
and find them to be conspecific. 


Heteristius cinctus (Osburn and Nichols) 
(Figure 16.) 


Dactyloscopus cinctus Osburn and Nichols, 1916: 176 (origi- 
nal description; Cape San Lucas, Mexico). 

Heteristius jalisconis Myers and Wade, 1946: 160 (original 
description; Jalisco, Mexico). 


DIAGNOSIS: Diagnostic characters are those of 
the genus. 


DESCRIPTION: Measurements (mm) of 27.9-mm- 
SL lectotype follow: depth of caudal peduncle 
1.5; body depth 4.1; predorsal length 5.7; pre- 
anal length 8.6; pelvic to anal distance 3.4; head 
length 6.1; head breadth 3.2; maxillary to upper 
opercular angle 5.7; maxillary to upper preoper- 
cular angle 3.9; eye diameter 1.5; postorbital 
length 3.4; preorbital length 1.0; maxillary length 
2.2. See Tables 2-8 for meristic data. 

Body broad and deep in front, compressed 
and slender caudad; dorsum of head somewhat 
rounded, lateral profile a descending arch from 
nape to tip of lower jaw; maximum head breadth 
near posterior margin of preopercle, not strongly 


156 


narrowed in front, lower jaw slightly protruding 
and rounded in dorsal profile (Fig. 5); lips with 
well developed simple fimbriae, most numerous 
below, frequency ontogenetic, at least on lower 
lip (Fig. 15; Table 10); eye dorsolateral, protrud- 
ing but not on long protrusile stalk and without 
flaps or papillae; anterior naris a short tube with 
distal valve-like emargination, located close be- 
hind anterior rim of preorbital; posterior naris a 
pore with slightly elevated margin, located be- 
hind anterior naris and adjacent to anterior rim 
of orbit; preopercle with a closely set series of 
about 12-14 canals, usually unbranched, each 
with a distal pore and often with a few smaller 
pores between this and canal base (Fig. 1); infra- 
orbital canal typically with single pores in pre- 
orbital branch and with two or more pores in 
lachrymal branch and at most posterior posi- 
tions; opercle with a fringe of up to 12. slender 
unbranched fimbriae overlying upper posterior 
margin, frequency ontogenetic (Fig. 7; Table 10); 
opercles broadly expanded, membranous mar- 
gins cover pectoral- and pelvic-fin bases and 
overlap on underside of head. 


Dorsal-fin origin on nape, in advance of upper 
opercular angle (Fig. 13); dorsal fin with a 2- 
spined anterior finlet, usually with one (42%) or 
two (56%) isolated spines between finlet and 
continuous dorsal fin; height of spinous portion 
of continuous dorsal fin nearly thrice that of 
finlet and about 25% less than longest seg- 
mented dorsal ray, the fin notched or emargin- 
ate between spines and segmented rays. Posteri- 
ormost dorsal and anal rays usually free, some- 
times bound to peduncle by fragile membranes; 
last anal ray reaches beyond rear of hypural, last 
dorsal ray somewhat shorter, caudal peduncle 
entire, without notch in ventral margin (Fig. 2); 
caudal fin rounded; segmented caudal-fin rays 
12, always simple, the upper and lowermost 
shorter than median rays. 


Pectoral fin-rays 14 in 79% of 208 counts, 6 
fish had 13 rays in each fin, whereas 10 had bi- 
lateral counts of 15; pectoral fin somewhat 
pointed, 4th-6th rays from dorsalmost the long- 
est, rays gradually shorter but not enlarged ven- 
trad; tip of straightened and adpressed fin 
reaches well beyond deflection of lateral-line 
arch, reaches vertical between dorsal-fin ele- 
ments 14 and 18, usually between 14 and 16; 
pelvic fin |, 3, tips of segmented rays free with 
emarginate membranes between, longest ray 
usually falls short of anus; all fin-rays simple in 
examined material. 


PROC. CALIF; ACAD. SCI. 4TH SER., VOL: 41;;NOme 


Lateral line continuous, originates above and 
in advance of upper opercular angle (on or just 
before vertical from 1st dorsal spine), arches up- 
ward to follow close to dorsal-fin base, deflects 
between verticals from dorsal spines 8-12 and 
continues midlaterally to terminate as the last 
scale overlying caudal-fin base (Fig. 2); canals of 
straight lateral-line scales generally parallel to 
longitudinal body axis, canal of last scale may be 
angled slightly ventrad; scales of upper part of 
lateral-line arch persistent, posterior margins 
somewhat trilobate; scales of descending arch 
and straight lateral line somewhat deciduous; 
canal of straight lateral-line scales without a ter- 
minal branch anteriad, usually with a dorsal pore 
near middle or anterior third of canal. Area 
above lateral-line arch completely scaled in late 
juveniles and adults (Fig. 13), usually with three 
scales between base of finlet and 1st lateral-line 
scale and one scale between apex of arch and 
dorsal-fin base; venter, between anus and pelvic- 
fin base, and sides (except for pectoral axil) com- 
pletely scaled; head, pectoral-fin base and all 
fins, except for caudal base, naked; 9 longitudi- 
nal scale rows cross vertical near origin of straight 
lateral line. 


Premaxillaries somewhat protractile, pedicels fall 
just short of rear margin of orbit; jaw teeth coni- 
cal, somewhat curved inward, about three rows 
near symphyses but in single row behind; vomer 
and palatines edentate; tongue narrowly round- 
ed, basihyal spatulate; three complete gill arches 
plus a hemibranch with pore behind; no rakers 
on 1st gill arch; pseudobranchiae 7-9 (3 exam- 
ined); branchiostegals 6; without predorsal 
bones; proximal pterygiophore of 1st dorsal 
spine broad, inserted in advance of 1st neural 
spine (Fig. 3); abdominal vertebrae 10 in radio- 
graphs; caudal fin with 3 spiniform elements 
above and below, upper and lower hypurals each 
bear 6 segmented rays (in 3 cleared specimens). 


Color in alcohol. —Ground color is light tan 
to pale, markings tan to dark brown. Dorsum of 
head capped by a mottled brownish blotch be- 
hind eyes followed by a narrow pale bar cross- 
ing between upper margins of preopercles; short 
streaks of brown radiate from eye and cross up- 
per and lower jaw near symphysis; underside of 
lower jaw often with two small blotches anteriad; 
remainder of head mainly pale. From above, dor- 
sum crossed by about 12 brown bars from nape 
to caudal base, bar length irregular, about equal 
to pale interspaces. In lateral aspect, sides crossed 
by three prominent bars and there is an elongate 


DAWSON: EASTERN PACIFIC SAND STARGAZERS 


blotch on caudal base; these markings persist in 
long-preserved and otherwise faded material. The 
anterior bar is located near origin of continuous 
dorsal fin and extends slightly below middle of 
side; 2nd bar (below 15th-19th dorsal spines) 
and 3rd bar (below 9th-12th segmented rays) ex- 
tend ventrad to anal-fin base. Between these 
characteristic markings, bars crossing dorsum ter- 
minate on 1st or 2nd scale below dorsal-fin base 
and there are short stripe-like markings on mid- 
line and just above anal-fin base; pale areas else- 
where often irregularly flecked with brown. Dor- 
sal finlet mainly brown, tip of 1st spine pale; 
remainder of dorsal fin usually pale, occasionally 
with some brown shading on basal portions of 
rays located over dark bars; middle of caudal fin 
crossed by faint band of micromelanophores, 
other fins mainly pale. 


REMARKS: The short and partly inaccurate de- 
scription of Dactyloscopus cinctus is based on 
three specimens, ‘‘about the same size,”’ collected 
23 March 1911. Neither figure nor description is 
specifically referred to the holotype which is 
numbered AMNH 5206. There are no specimens 
now shelved under this number at the American 
Museum of Natural History, but there is a lot of 
three fish (AMNH 5394) with an external jar label 
bearing the pencil notation, ‘‘Type and paratype.” 
The jar contains a pencil note stating, ‘‘Cape San 
Lucas Mch. 23/11’’ and a printed ALBATROSS 
label penciled, ‘““Dactyloscopus cinctus N............ 
INNAGGM Zoe ce eee oxen. Cape San Lucas.’ | am advised 
by J. E. Darovec, Jr. (AMNH) that catalog cards 
for AMNH 5206 and 5394 show the same data, 
and it appears that these numbers were intended 
to accommodate the three specimens mentioned 
by Osburn and Nichols (1916). It is not known 
whether these authors actually separated the 
selected holotype from the two remaining fish 
or whether the holotype was subsequently re- 
united with AMNH 5394, but there is little doubt 
that the three original specimens are now in- 
cluded in this single lot. In any event, it is now 
impossible to identify the originally intended 
holotype and | treat these fish as syntypes. | 
select the best preserved specimen (27.9 mm SL) 
as the lectotype of Dactyloscopus cinctus. This 
fish (AMNH 5206) has about 27 fimbriae on the 
lower lip, about 21 scales in lateral-line arch, 
scaled venter and the characteristic body bars 
are distinct on both sides (for additional data, 
see measurements above and Tables 2, 3 and 7). 
The two remaining specimens (29 mm SL) are 
paralectotypes (AMNH 5394). 


1!57/ 


Labial fimbriae are distinct and characteristic 
body bars are present at 14 mm SL; squamation 
is apparently complete in some 16-mm fish. No 
specimens were found with modified anterior 
anal-fin rays or enlarged pectoral fins that char- 
acterize sexually mature (transformed) males of 
other sand stargazers. Examination of radio- 
graphs showed no evidence of the broadened 
anterior anal pterygiophores commonly associ- 
ated with the externally modified anal rays. A 
number of ovigerous females were present 
among examined material and it seems unlikely 
that mature males were not included. No further 
efforts were made toward sex determination, but 
it is highly probable that anal rays and pectoral 
fins are not modified in Heteristius cinctus. Ab- 
sence of such modifications would suggest that 
adult males do not carry incubating egg clusters 
beneath the pectoral fins as reported for other 
dactyloscopids (BOhlke and Chaplin, 1968; Petti, 
1969). The complete squamation of venter and 
adjacent sides, naked or largely so in other dac- 
tyloscopids, would seem to support this assump- 
tion. Among material examined, the smallest 
ovigerous female is 22 mm SL; a 32-mm fish con- 
tained 171 developing eggs in the right gonad. 

| have collected specimens at depths of less 
than a meter, others have been taken with 
SCUBA in 15 m and maximum recorded depth is 
from a dredge sample in 27.4m. There are no 
collections from low salinity or estuarine habitats. 
Heteristius cinctus is known from Bahia Magdal- 
ena to the vicinity of Punta Arena on the Baja 
California peninsula. It is apparently absent from 
the Gulf of California but reappears in collections 
from a number of insular and mainland shores 
from Nayarit, Mexico south to Cabo San Fran- 
cisco, Ecuador (Fig. 8). 


MATERIAL EXAMINED: Three hundred fifty-eight specimens, 
14-40 mm SL, including lectotype and two paralectotypes. 

Lectotype. —AMNH 5206 (27.9 mm SL), Mexico, Baja Cali- 
fornia Sur, Cape San Lucas, seine, 28 March 1911, Town- 
send ALBATROSS Expdn. 

Paralectotypes. —AMNH 5394 (2, 29 mm SL), taken with 
lectotype. 

Other Material. —MEXICO, Baja California Sur: CAS 32041 
and LACM 31174-46; SIO 61-248, 62-706, 62-720, 64-42 and 
64-43; UCLA W59-248. —Nayarit: GCRL 8500 and 13363; 
UCLA W58-5 and W58-8. —Islas Tres Marias: GCRL.13341, 
13362 and SIO 62-25. —Jalisco: LACM 23185 (holotype of 
H. jalisconis). —Oaxaca: LACM 23186 and CAS-SU 39961 
(paratypes of H. jalisconis). 

COSTA RICA, Puntarenas: GCRL 11813. —Isla del Cano: 
GCRL 13360 and 13361; LACM 32546-47, 32549-59 and UCR 
674-5. 


PANAMA, Isla Coiba: SIO 71-211 and 71-213. 
ECUADOR: USNM 116221. 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 2 


158 


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DAWSON: EASTERN PACIFIC SAND STARGAZERS 


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160 


ACKNOWLEDGMENTS 


Appreciation is expressed to curators of the 
various repositories for the loan of specimens 
and other courtesies. Special thanks are due 
W. A. Bussing (UCR), R. J. Lavenberg and C. 
Swift (LACM), R. H. Rosenblatt (SIO) and B. W. 
Walker (UCLA) for making their collections avail- 
able for study. Miss Pearl Sonoda (CAS) provided 
considerable help with problems concerning 
CAS-SU collections. For permission to work in 
Chile and for assistance during my visit, | thank 
N. Bahamonde, F. R. Manns, G. Pequeno-R., G. 
Revuelta and J. Tomicic. Drs. J. E. Bohlke (ANSP) 
and V. G. Springer (USNM) provided useful com- 
ment and suggestions on portions of this work. 
C. L. Smith (AMNH) conducted the search for 
data on the type of Dactyloscopus cinctus. 

Drawings are by Anne Langenfeld; computer 
services were provided by Paul Poole and Mrs. 
Lois Coquet; F. N. Jackson is thanked for general 
laboratory assistance and radiographs. Special 
acknowledgment is due Mrs. Betty Heal for ex- 
pert secretarial assistance and for her interest in 
this study and others now in progress. This work 
was supported in part by National Science Foun- 
dation Grant GB-31053X. 


LITERATURE CITED 


Bohlke, J. E. 1968. The descriptions of three new sand star- 
gazers (Dactyloscopidae) from the tropical west Atlantic. 
Notulae Naturae 414: 1-16. 

Bohlke, J. E. and C. C. G. Chaplin. 1968. Fishes of the 
Bahamas and adjacent tropical waters. Livingston Publ. 
Co., Wynnewood, Pa. 771 pp. 

Dawson, C. E. 1969. A new eastern Pacific sand stargazer 
(Pisces: Dactyloscopidae). Copeia 1969(1): 44-51. 


PROG. GALIF; AGAD. SGI 47H SER VOL. 41 NOme 


1974. Studies on eastern Pacific sand stargazers 
(Pisces: Dactyloscopidae). 1. Platygillellus new genus, 
with descriptions of new species. Copeia 1974(1): 39-55. 

1975. Studies on eastern Pacific sand stargazers 
(Pisces: Dactyloscopidae). 2. Genus Dactyloscopus, with 
description of new species and subspecies. Natural His- 
tory Museum, Los Angeles County, Science Bulletin 22: 
1-61. 


. 1976. Studies on eastern Pacific sand stargazers. 3. 


Dactylagnus and Myxodagnus, with description of a new 
species and subspecies. Copeia 1976(1): 13-43. 

Fowler, H. W. and B. A. Bean. 1923. Descriptions of eigh- 
teen new species of fishes from the Wilkes Exploring 
Expedition, preserved in the United States National 
Museum. Proceedings of the United States National 
Museum 63: 1-27. 

Gilbert, C. H. 1890. Scientific results of explorations by the 
U.S. Fish Commission Steamer ALBATROSS. No. XII. 
—A preliminary report on the fishes collected by the 
steamer ALBATROSS on the Pacific coast of North 
America during the year 1889, with descriptions of 
twelve new genera and ninety-two new species. Pro- 
ceedings of the United States National Museum 13: 


49-126. 
. 1892. Scientific results of explorations by the U.S. 
Fish Commission Steamer ALBATROSS. No. XXII. —De- 


scriptions of thirty-four new species of fishes collected 
in 1888 and 1889, principally among the Santa Barbara 
Islands and in the Gulf of California. Proceedings of 
the United States National Museum 14: 539-566. 

Jordan, D. S. 1897. Notes on fishes, little known or new 
to science. Proceedings of the California Academy of 
Sciences, ser. 2, 6: 201-244. 

Myers, G. S. and C. B. Wade. 1946. New fishes of the famil- 
ies Dactyloscopidae, Microdesmidae, and Antennariidae 
from the west coast of Mexico and the Galapagos Is- 
lands. Allen Hancock Pacific Expeditions 9: 151-178. 

Osburn, R. C. and J. T. Nichols. 1916. Shore fishes collected 
by the ‘Albatross’ Expedition in Lower California with 
descriptions of new species. Bulletin of the American 
Museum of Natural History 35(16): 139-181. 

Petti, J. C. 1969. Behavioral and morphological adaptations 
to burrowing of two species of dactyloscopid fishes 
from the northern Gulf of California. Unpbl. Master’s 
thesis, University of Arizona, Tucson. 


OF THE 


zi PROCEEDINGS 


LIBRANN 


sa 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 3, pp. 161-168, 5 figs., 1 table. 


April 15, 1977 


THREE NEW SPECIES OF INDO-PACIFIC MORAY EELS 
(PISCES: MURAENIDAE) 


John E. McCosker 
Steinhart Aquarium, California Academy of Sciences, 
Golden Gate Park, San Francisco, California 94118 


John E. Randall 


Bernice P. Bishop Museum, Honolulu, Hawaii 96818 


ABSTRACT: Three new species of Indo-Pacific muraenid eels are described and illustrated: Gymnothorax breedeni 
n.sp. an eel from moderate depth reefs, from the Comoro, Amirante, Maldive, Marquesas, and Line islands; Uro- 
pterygius kamar n.sp. a deep-reef, burrowing eel from the Comoro, Amirante, Society, Solomon, Caroline, and 
Pitcairn islands; and U. goslinei n.sp. a shallow-reef eel from New Guinea, and the Caroline and Solomon islands. 


INTRODUCTION 


Recent deep water collections using rotenone 
ichthyocides in the tropical Indian and Pacific 
oceans have resulted in the discovery of two new 
moray eels. With a grant from the Charline 
Breeden Foundation, the senior author was able 
to make the first extensive series of reef fish col- 
lections from the remote Archipel des Comores 
(Comoro Islands). Those specimens complement 
the extensive collections made by the junior 


author throughout Oceania, and extend the range 
of many Pacific species to include the western 
Indian Ocean. In the process of describing these 
morays, a third and distinctive new species was 
discovered among the collections of reef fishes 
made by the George Vanderbilt Foundation, now 
housed at the California Academy of Sciences. 
Measurements are straight-line, made either 
with a 300-mm ruler with 0.5-mm gradations (for 
total length, trunk length, and tail length) and re- 
corded to the nearest 0.5 mm, or with dial cali- 


CALIFORNIA ACADEMY OF SCIENCES / GOLDEN PATE PARK / SAN FRANCISCO 


162 


pers (all other measurements) and recorded to 
the nearest 0.1 mm. Body length comprises head 
and trunk lengths. Head length is measured from 
the snout tip to the posterodorsal margin of the 
gill opening; trunk length is taken from the end 
of the head to mid-anus; maximum body depth 
does not include the median fins. Vertebral 
counts (which include the hypural) were taken 
from radiographs. Materials used in this study 
are housed at the following institutions: Austral- 
ian Museum, Sydney (AMS); Academy of Natural 
Sciences of Philadelphia (ANSP); Bernice P. 
Bishop Museum (BPBM); California Academy of 
Sciences (CAS, including George Vanderbilt 
Foundation Collections, GVF); U.S. National Mu- 
seum of Natural History (USNM); J.L.B. Smith In- 
stitute of Ichthyology, Rhodes University (RUSI); 
and the Scripps Institution of Oceanography 
(SIO). Paratypes of Uropterygius kamar will be 
deposited at the Muséum National d'Histoire 
Naturelle Paris (MNHN), and the British Museum 
(Natural History). 


ACKNOWLEDGMENTS 


We wish to thank the following individuals: 
Maurice C. Giles for photographic assistance; 
William C. Ruark and James E. Gordon for the 
preparation of radiographs; Virginia M. Gregory 
(Figs 3 a-c), Katherine P. Smith (Fig. 5), and 
Mary H. Fuges (Figs. 1-2) for their artwork; Lillian 
J. Dempster and William N. Eschmeyer for their 


PROG) GAIA AGAD SS Gir. 4ltieS ERs eV © LertulaN © es 


critical reading of this manuscript; and the cur- 
ators of fish collections for allowing us to exam- 
ine specimens. James E. Bohlke kindly made 
available his specimens of the new species and 
the illustrations prepared by Mary Fuges. Ran- 
dall’s collections were made possible in part by 
grants from the National Geographic Society and 
the American Philosophical Society. A portion 
of McCosker’s work was supported by funds 
from the Charline Breeden Foundation. We are 
especially grateful to the people of the Archipel 
des Comores, and in particular, to Chief Moham- 
med Ali Chabane and Prince Nacr-Ed-Dine, for 
allowing us to collect specimens. 


TAXONOMY 


Gymnothorax breedeni McCosker & Randall, new 
species 
(Figures 1-3, Table 1.) 


Gymnothorax sp.: Randall & Helfman, 1972: Figs. 5-6 (photo- 
graphs of Marquesas specimens mentioned in text). 

Gymnothorax sp.: McCosker & McCosker, 1976: 26 (Comoran 
specimens mentioned in text). 


MATERIAL EXAMINED. Holotype: CAS 35250, 530 mm. Archi- 
pel des Comores, Isle Grande Comore, Itsandra, 0.5 km 
north of Hotel Itsandra. Captured in 20 m among coral and 
rock within a large cave, 50 m offshore, on 19 February 1975 
by J. —. McCosker, S. McCosker, M. D. Lagios, L. Gunther, 
and D. C. Powell, using Pronoxfish ichthyocide. Paratypes: 
USNM 215283 (formerly BPBM 7763), 645 mm, Line Is., 
Washington Id., 15 m, J. E. Randall. BPBM 11903, 524 mm, 
Marquesas Is., Tahuata, 10 m, J. E. Randall. ANSP 134222, 
496 mm, Amirantes Is., D’Arros Id., 4 m, J. E. Bohlke. 


TABLE 1. Counts, and proportions in thousandths, of the holotypes and paratypes of Gymnothorax breedeni (3 paratypes), 
Uropterygius goslinei (7 paratypes), and U. kamar (9 paratypes). TL = total length, HL = head length. 
‘ie i G. breedeni Wnaestine! ; : U. bower “— 

Holotype Mean Range Holotype Mean Range Holotype Mean Range 

Total length 530 mm 549 496-645 483 mm_ 271 149-527 249 mm 232 164-345 
Vertebrae 128 128.5 128-129 127, 127.9 124-130 13h 136.3 134-138 
Head/TL 152 130 122-136 116 116 105-121 102 103 95-108 
Trunk/TL 340 336 302-356 300 309 = 300-333 305 306 299-307 
Tail/TL 528 533 512-562 584 574 547-587 592 SZ 571-606 
Snout/HL 181 HS) 171-186 161 170 161-197 161 158 132-183 
Jaw/HL 379 374 353-398 368 386 328-458 369 360 8279-423 
Eye/HL 79 87 76-112 73 65 55- 76 U5) 69 63- 77 
Interorbital/ HL 119 140 119-170 AS 11 105-133 110 97 79-119 
Depth/HL 543 622 475-765 482 405 348-482 578 353 316-402 
Width/HL 314 364 312-470 359 264 205-339 267 224 195-267 


McCOSKER & RANDALL: NEW MORAY EELS 


163 


Figure 1. Gymnothorax breedeni, paratype, ANSP 134222, 496 mm. 


DIAGNOSIS. A moderate length species of Gym- 
nothorax with anus before mid-body, tubular an- 
terior nostrils and slightly raised posterior nos- 
trils, uniserial jaw teeth, vomerine teeth either 
absent or few, and a brown coloration with dis- 
tinctive black areas behind the eye, at the rictus, 
and over the gill opening. 


DESCRIPTION OF HOLOTYPE. (Counts and pro- 
portions of the holotype and the three paratypes 
are given in Table 1.) Greatest depth of body 14 
times in total length (TL). Body width 1.7 times 
in depth. Tail longer than body, distance from 
snout to anus 2.12 times in TL. Head 7.6 times 
and trunk 2.9 times in TL. Dorsal fin origin ahead 
of gill opening, the predorsal distance ca. 8.8 
times in TL. Dorsal fin height at gill opening ca. 
3.5 times in body depth. Snout 5.5 times and 
upper jaw 2.6 times in head length (HL). Eye 12.7 
times in HL and 2.3 times in snout, closer to 
rictus than tip of snout. Fleshy interorbital width 
8.4 times in head. Gill opening nearly horizontal, 
its center slightly below mid-body, its length 
about equal to diameter of eye. 

Anterior nostril tubular, minute, not reaching 
snout tip. Posterior nostril well ahead of eye, a 
vertical line drawn from it falls nearly 0.5 eye 
diameter in front of eye; opening small, nearly 


circular, forming a small tube in larger individ- 
uals. 


Jaws subequal, the mouth closing completely. 
Teeth in jaws (Fig. 3) uniserial, nearly subequal, 
stout, pointed, and finely serrated. Two depres- 
sible long canines along mid-line of intermaxil- 
lary, the posterior the largest tooth in the mouth. 
Three pairs of depressible canines behind man- 
dibular symphysis. 

Number of vertebrae 128, 52 before anal fin. 
The first dorsal pterygiophore arises above the 
third vertebra. 


Head pores (Fig. 2) present but not obvious. A 
single pore anterior and proximal to, and a sec- 
ond pore below the base of the anterior nostril. 
Six pores along the mandible, the second through 
fifth the largest. Four equally spaced pores along 
the upper jaw, the first beneath the nostril base, 
the last beneath the center of the eye. A single 
pore between each anterior and posterior nostril. 

Color in isopropyl alcohol brown overlain with 
a darker speckling throughout, although slightly 
lighter along the belly. Anus and gill openings 
within black areas. A prominent black spot. be- 
hind each eye, with a black slash beginning at 
the ventral margin of the orbit and ending about 
an eye’s diameter behind the rictus. A black spot 


164 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 3 


ey aS 
eh 


ENN es 
MSGEe Mae Sep ete 


Figure 2. Gymnothorax breedeni, paratype, ANSP 134222, 496 mm. 


smaller than the eye present beneath the rictus 
of some specimens. Margin of caudal cream, 
yellowish in life. 


ETYMOLOGY. Named in honor of Vic E. Breeden, 
president of the Charline H. Breeden Foundation 
which made this collection and study of Comoran 
fishes possible. 


REMARKS. The closest relative of Gymnothorax 
breedeni appears to be G. flavimarginatus (Rup- 
pell, 1828), a wide-ranging species which was 
either observed or collected at each of the locali- 
ties where we have taken the new species. G. 
breedeni differs from G. flavimarginatus in_ its 
coloration (viz., the black markings associated 
with the eye and jaw and the restriction of the 
light fin edging to the tail) and in its reduced or 
lack of vomerine dentition. 


The new species is similar in appearance to 
G. monostigma (Regan, 1909) in having a black 
blotch behind the eye. G. monostigma is readily 
separable in having the lower jaw pores and pos- 
terior nostril lying within white spots and in lack- 
ing a black area around the gill opening. 


The posterior nostril of the new species be- 
comes more tubular with age (see Fig. 2). This 
condition occurs among several species of vari- 
Ous moray genera and is considered by us not 
significant as a generic character. 


A color photograph which nicely illustrates the 
facial coloration of living G. breedeni appeared 
in Randall and Helfman (1972). Besides the typi- 
cal material herein described, another specimen 
of this distinctive species was observed but not 
collected by Randall at Villingili Island, Maldives, 
at about 10 m depth. 


Uropterygius kamar McCosker & Randall, new 
species 
(Figures 3b & 4, Table 1.) 


MATERIAL EXAMINED. Holotype: CAS 35251, 249 mm. Archi- 
pel des Comores, Isle Grande Comore, Itsandra, 0.5 km 
north of Hotel Itsandra. Captured in 25 m over coral sand 
slope, 50 m offshore, using Pronoxfish ichthyocide, on 19 
February 1975 by J. E. McCosker, S. McCosker, M.D. Lagios, 
L. Gunther, and D. C. Powell. Paratypes: CAS 35252, 20(73- 
224 mm), and RUSI 4557, 5(112-215 mm), collected with the 
holotype. Collected by J. E. Randall using ichthyocides or a 
hand spear, generally associated with a coral rubble bottom 
between depths of 3 to 55 m: BPBM 12051, 345 mm, Tahiti, 
Popote Bay; BPBM 9442, 2(211-228 mm), Palau Is., Augelpela 
Reef; USNM 215281 (formerly BPBM 15665), 219 mm, Solo- 
mon ls., Florida Id.; BPBM 16471, 4(150-325 mm), Pitcairn 
Id., Oeno Id.; BPBM 16606, 297 mm, Pitcairn Id., Oeno Id.; 
BPBM 17005, 136 mm, Pitcairn Id.; BPBM 17047, 327 mm, 
Pitcairn Id. Collected by J. E. Bohlke et al.: ANSP 124794, 
3(145-198), Amirantes Is., St. Joseph Id., 15-27 m. ANSP 
134217, 11(124-365), Amirantes Is., D’Arros Id., 20-30 m. 


DIAGNOSIS. A small species of Uropterygius 
with vertical fins restricted to the tail tip, anus 
before mid-body, two prominent cephalic lateral 


McCOSKER & RANDALL: NEW MORAY EELS 


165 


2 2 Aas TR 
WAN oe b Whe SAdTALUN ISH 


"RABE 
a, 
SSS 


AS \se ea 


BLE 


=, 


ALA 


CLPYIP PVA IPP I 2p EEE 


S 


5 


Figure 3. Dentition of: (A) holotype of Gymnothorax breedeni, CAS 35250, 530 mm; (B) holotype of Uroptery- 
gius kamar, CAS 35251, 249 mm; and (C) paratype of Uropterygius goslinei, ANSP 117434, 527 mm. 


lines pores, posterior nostril not closely associ- 
ated with a supraorbital pore, triserial maxillary 
dentition, white elongate anterior nostrils, and 
a prominent white band in front of the eye. 


DESCRIPTION OF HOLOTYPE. (Counts and pro- 
portions of the holotype and nine paratypes are 
given in Table 1.) Body subrectangular in cross 
section, laterally compressed throughout, _ its 
greatest depth 26 times in total length (TL). Body 
width 1.4 times in depth. Tail longer than body, 
distance from snout to anus 2.45 times in TL. 
Head 9.8 times and trunk 3.3 times in TL. Dorsal 
fin restricted to tail, arising about one-half head 
length (HL) from tail tip. Snout 6.2 times and 


upper jaw 2.7 times in HL. Eye 13 times in HL 
and 2.2 times in snout, equidistant between 
snout and rictus. Fleshy interorbital width 9.1 
times in HL. Gill opening small, ovate and hori- 
zontal, lying about mid-body, its length less than 
eye diameter. 

Anterior nostrils elongate, tubular, nearly equal 
in length to eye, extending noticeably beyond 
snout. Posterior nostril within the interorbital 
space, above and slightly in advance of middle 
of eye, lying within a short tube, the proximal 
edge flush with the head. 

Jaws subequal, the mouth closing completely. 
Teeth in jaws (Fig. 3b) pointed, recurved. Upper 
jaw teeth triserial, the outer row teeth small, 


166 


PROG (GALEIEVAGAD. SGlF 4a SERS VOL 41 NOs 


Figure 4. Head and anterior trunk region of paratypes of Uropterygius kamar. A. Uniform coloration, BPBM 
9442, 228 mm. B. Mottled coloration, BPBM 12051, 345 mm. 


numerous and close-set, the middle row teeth 
somewhat irregular and intermediate in size, the 
inner row teeth the largest, comprised of widely- 
spaced depressible fangs. Lower jaw teeth bi- 
serial, except triserial anteriorly, the outer row 
teeth small, numerous and close-set, the inner 
row teeth widely-spaced, depressible fangs, larg- 
est anteriorly. A series of depressible fangs in 
the intermaxillary region, separated from the 
uniserial vomerine teeth by a gap. 


Number of vertebrae 137, 49 before anal 
opening. Median fins restricted to the end of 
the tail, appearing above the 115th vertebra on a 
radiograph. 

Head pores distinct, usually associated with 
white markings along snout and lips. Anterior 
nostril bases bordered laterally and ventrally by 
pores. Six pore pairs along upper and lower 
jaws, alternating in position. Two  supraorbital 
canal pores. Two cephalic lateral line pores, fol- 
lowed by a minute series of lateral line pores 
passing along the mid-body to the tail tip. 

Color in isopropyl alcohol tan to chocolate 
brown, distinctively marked by a variable series 
of white bands along the chin, throat, snout and 


nape. A prominent band passes obliquely from 
the anterior edge of the eye through the poster- 
ior nostril and meets its partner along the inter- 
orbital space. Anterior nostrils white and con- 
nected between their bases by a white stripe. 
Inside of mouth white. The trunk of small speci- 
mens often has a faint series of lighter, broken 
bands. 


ETYMOLOGY. Named in honor of the Archipel 
des Comores, the location of capture of many 
of the type specimens. The newly independent 
Comoran Republic has derived its name from 
kamar, an Arab word for the moon, in reference 
to the moon-like quality of the volcanic surface 
of those lovely islands. 


REMARKS. This new species is most similar to 
Uropterygius xanthopterus Bleeker (1859), from 
which it is easily separable on the basis of its 
distinctive coloration. U. kamar also differs from 
specimens we have identified as U. xanthopterus 
(CAS 35254 from Kapingamarangi and SIO 73-205 
from Australia) in having more vertebrae and a 
more anterior median fin origin (as viewed from 
radiographs). The new species is also similar to 


McCOSKER & RANDALL: NEW MORAY EELS 


167 


Figure 5. Head and mid-trunk region of holotype of Uropterygius goslinei, CAS 35253, 483 mm. 


U. alboguttatus Smith (1962), described from the 
western Indian Ocean, which is more closely 
related to U. xanthopterus (and possibly synony- 
mous) and differs markedly in its coloration. U. 
kamar differs from its other congeners particu- 
larly in its coloration, multiserial dentition, and 
the presence of two cephalic lateral line pores. 


Uropterygius goslinei McCosker & Randall, new 
species 
(Figures 3c & 5, Table 1.) 


MATERIAL EXAMINED. Holotype: CAS 35253, 483 mm. Palau 
Is., Kayangel Atoll. Captured in a lagoon 200 m east of the 
west side of the atoll, in 0-3 m depth over a sand and coral- 
line algal bed with coral heads, using Noxfish ichthyocide by 
R. Gaines and party on 8 October 1956. Paratypes: Collected 
in shallow water over sand and coral bottoms using rotenone 
ichthyocides. CAS 13972, 212 mm, and BPBM 19927, 149 mm, 
Kapingamarangi Atoll. CAS 29103, 213 mm, Papua, New 
Guinea, Egum Atoll. USNM 215282 (formerly GVF-1955, Sta. 
25), 151 mm, Palau Is., Koror Id. AMS.1. 17600-001 (formerly 
CAS 13973), 252 mm, Palau Is., Kayangel Atoll. ANSP 117434, 
527 mm, Solomon Is., Bougainville. ANSP 124812, 270 mm, 
Papua, New Guinea, Madang Harbor. 


DIAGNOSIS. A moderate length species of Uro- 
pterygius with vertical fins restricted to the tail 
tip, anus before mid-body, two cephalic lateral 
line pores, posterior nostril not closely associated 
with a supraorbital pore, short jaws, triserial jaw 
teeth, and a tan coloration overlain by narrow 


black bands and white spots and two distinct 
black stripes behind the eye. 


DESCRIPTION OF HOLOTYPE. (Counts and pro- 
portions of the holotype and seven paratypes are 
given in Table 1.) Body subrectangular in cross 
section, laterally compressed throughout, _ its 
greatest depth 18 times in total length (TL). Body 
width 2.5 times in depth. Tail longer than body, 
the distance from snout to mid-anus 2.4 times in 
TL. Head 8.6 times and trunk 3.3 times in TL. 
Dorsal fin restricted to tail. Snout 6.2 times and 
upper jaw 2.6 times in head length (HL). Eye 14 
times in HL and 2.2 times in snout, its center 
slightly in advance of middle of jaw. Fleshy in- 
terorbital width 8.7 times in HL. Gill opening 
small, ovate and horizontal, lying about mid- 
body, its length less than eye diameter. 

Anterior nostril tubular, much smaller than 
eye, not extending beyond snout. Posterior nos- 
tril within the interorbital space, above and in 
advance of eye, lying within a short tube (Fig. 5). 

Jaws subequal, the mouth closing completely. 
Teeth in jaws (Fig. 3c) pointed, recurved. Upper 
jaw teeth triserial, the outer row teeth small, 
numerous and close-set, the middle row teeth 
intermediate in size, the inner row teeth the 
largest, comprised of depressible fangs. Lower jaw 
teeth triserial, the outer row teeth small, num- 


168 


erous and close-set, the middle row teeth inter- 
mediate and not extending beyond mid-jaw, the 
inner row teeth the largest, comprised of de- 
pressible fangs. Intermaxillary region is crowded 
by a group of large depressible fangs, separated 
from the uniserial vomerine teeth by a gap. 

Number of vertebrae 127, 48 before anal open- 
ing. Median fins restricted to end of tail, above 
the 114th vertebra on a radiograph. 

Head pores indistinct. Anterior nostril bases 
bordered laterally and ventrally by pores. Five 
pore pairs along upper and lower jaws. Two 
supraorbital canal pores. Two cephalic lateral 
line pores, followed by a minute series of lateral 
line pores passing along the mid-body to the tail 
tip. 

Color in isopropyl alcohol cinnamon, darker 
dorsally, overlain with narrow black lines which 
are broken along the lateral mid-line, with white 
punctations irregularly located between the lines. 
Two black lines originate at the posterior dorsal 
and ventral margins of the eyeball and extend 
2-3 eye lengths posteriorly. 


ETYMOLOGY. Named in honor of William 
Alonzo Gosline in recognition of his numerous 
contributions to the study of apodal fishes. 


REMARKS. Following Gosline’s (1958) key to the 
central Pacific species of Uropterygius, the new 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 3 


species would be identified as U. marmoratus 
(Laceépede, 1803). U. goslinei differs in having 
two rather than a single cephalic lateral line pore 
and in its coloration. The new species is similar 
to U. xanthopterus and U. kamar in having two 
pores, but differs in its coloration and its com- 
plex dentition, and in attaining a larger size. 


LITERATURE CITED 


Bleeker, Pieter. 1859. Over eenige vischsoorten van de Zuid- 
kustwateren van Java. Nat. Tijdschr. Neder.-Indie, 19: 
329-352. 

Gosline, William A. 1958. Central Pacific eels of the genus 
Uropterygius, with the descriptions of two new species. 
Pacific Sci., 12(3):221-228. 

Lacépede, (Comte) B. G. E. 1803. Histoire naturelle des pois- 
sons....deédiée au citoyen Lacépede. Vol. 5, Paris. 

McCosker, Sandra, and John £. McCosker. 1976. To the 
islands of the moon. Pacific Discovery, 29(1):19-28. 

Randall, John E., and Gene Helfman. 1972. Diproctacanthus 
xanthurus, a cleaner wrasse from the Palau Islands, with 
notes on other cleaning fishes. Trop. Fish Hobbyist, 
20(11):87-95. 

Regan, C. Tate. 1909. Descriptions of new marine fishes from 
Australia and the Pacific Ann. Mag. Nat. Hist., ser. 8, 
4:438-440. 

Ruppell, Wilhelm P. E. S. 1828. Atlas zu der Reise im nord- 
lichen Afrika. Zoologie. Fische des Rothen Meeres. 4 
vols. Frankfurt-a-M., 1826-1828. 

Smith, J. L. B. 1962. The moray eels of the western Indian 
Ocean and the Red Sea. Ichthyological Bulletin, Rhodes 
University, 23:421-444. 


NH PROCEEDINGS 
OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 4, pp. 169-182, 3 figs., 1 table. May 31, 1977 


TIGER BEETLES OF THE GENUS C/C/INDELA IN THE SULPHUR SPRINGS 
VALLEY, ARIZONA, WITH DESCRIPTIONS OF THREE NEW SUBSPECIES 
(CICINDELIDAE — COLEOPTERA) 


by 
Norman L. Rumpp 


3446 Bahia Blanca West 
Laguna Hills, California 92653 


ABSTRACT: The Sulphur Springs Valley of southeastern Arizona contains a large tiger beetle 
fauna. Seventeen species of Cicindela (Cicindelidae - Coleoptera) are reported; of these, 
three are described as new subspecies: C. willistoni sulfontis, C. pulchra dorothea and C. 
nevadica citata. Ecological associations of each species are detailed and a key is provided for 
identification. The relationships with other subspecies are mentioned, with special emphasis 
on the three subspecies described. The Cicindela fauna of the valley is shown to be related, 
for the most part, to the fauna of the mid-Rio Grande River System, indicating that past 
physical geology of the lower Southwest allowed hydrographic flow in an eastward direction, 
at least as early as the Miocene. In late Pliocene or early Pleistocene, stream flow began going 
in a westerly direction allowing some migration of species in that direction. This had a 
limited effect on the tiger beetles of the Sulphur Springs Valley because this valley remained 
isolated during most of the Pleistocene. 


INTRODUCTION 


Many of the western United States tiger 
beetles of the genus Cicindela have undergone 
subspeciation. This is particularly evident in the 
Southwest where profound geological changes 
have occurred, especially during late Cenozoic. 
These changes have produced environmental dis- 
locations and have altered climate in such di- 
verse ways as to induce strong ecological pres- 
sures on the various populations. The Sulphur 
Springs Valley of southeastern Arizona has been 


totally isolated geologically from adjacent valleys 
at least through the latter part of the Pleistocene 
when it became an area of internal drainage. 
The Cicindela fauna of this valley contains a large 
number of species, some of which have uniquely 
subspeciated. The subspecies described in this 
paper are isolated from the main populations of 
their respective species, some by a considerable 
distance, thus extending our knowledge of the 
geographical limits of these species. Students 
and collectors of tiger beetles have been at- 
tracted to this valley for the past twenty years be- 


CALIFORNIA ACADEMY OF SCIENCES/GOLDEN GATE PARK/SAN FRANCISCO 


170 


cause of its rich fauna. During this same period 
the author has conducted fifteen field trips into 
the valley, and has accumulated over two thou- 
sand specimens for this study. 


ACKNOWLEDGMENTS 


The author is grateful for the assistance given 
by many persons who made it possible to dis- 
cover and accumulate specimens for this paper. 
These include: Dr. Mont A. Cazier of Arizona 
State University, Tempe, Arizona; Mrs. Patricia 
Vaurie of the American Museum of Natural His- 
tory, New York, N.Y.; Dr. Oscar A. Cartwright of 
the U.S. National Museum, Washington, D.C.; 
the Reverend Bernard Rotger of Pagosa Springs, 
Colorado; Mr. Douglas K. Duncan of Globe, 
Arizona and Mr. Vernon G. Clifford of Graham, 
Washington who sent specimens from their pri- 
vate collections; Mr. Jay M. Sheppard while a 
student at California State University, Long Beach, 
Dr. John Stamatov of Armonk, New York, Dr. 
deWendler-Funaro of Williston Park, New York, 
and Dr. David L. Pearson of Pennsylvania State 
University, who participated in the field; finally, 
and most importantly, my late wife Dorothy who 
shared enthusiastically in many field trips to col- 
lect specimens and take notes. 


PHYSIOGRAPHY OF THE SULPHUR 
SPRINGS VALLEY 


The Sulphur Springs Valley lies in the moun- 
tainous region of Arizona, south of the plateau 
region, and east of the desert region. The moun- 
tains are of the block-fault type, similar to those 
of the Basin and Range province of Utah and 
Nevada, but appear to be older. The valley lies 
across the center of Cochise County and extends 
northward into the southern portion of Graham 
County; it trends in a north-northwestward di- 
rection from the Mexican border for about 145 
kilometers, with an average width of about 32 
kilometers. The valley covers about 4700 square 
kilometers and drains another 2600 square kilo- 
meters of the bordering watershed. The 32nd 
parallel crosses the valley just south of the large 
central playa, while the 110th meridian traverses 
to the west of the playa. The valley is bounded 
on both sides by mountain ranges which sepa- 
rate it on the east from the San Simon Valley, 
and on the west from the San Pedro Valley; both 
of these valleys drain northward into the Gila 
River basin. The Sulphur Springs Valley is sepa- 


PROG. GALIF. AGAD:* SGI-, 41H SER:;, VOI AT) INO 34 


rated on the southeast from the San Bernardino 
Valley by the Perilla Mountains, however, this 
smaller valley and the southern third of the 
Sulphur Springs Valley drain to the south into the 
Yaqui River system of Sonora, Mexico. 

The mountains that enclose the valley from the 
east include the Pinalino, Dos Cabezas, Chiri- 
cahua, Pedregosa, and Perilla mountains. The 
western chain includes the Galiuro, Winchester, 
Little Dragoon, Dragoon, and Mule mountains. 
The mean elevation of the valley is slightly over 
1300 meters. The climate is semi-arid as the cen- 
tral portion receives an average yearly rainfall of 
less than 30 centimeters. There is a rainy season 
which begins in mid-July and lasts through Sep- 
tember. 

The center of the valley contains a_ large, 
roughly triangular playa that covers about 132 
square kilometers and drains the upper two thirds 
of the valley. This playa lies a few kilometers 
south of the town of Willcox, at an elevation of 
1257 meters. During pluvial times this basin of 
internal drainage contained a lake that was ap- 
proximately 32 kilometers long by 18 kilometers 
wide, thus covering an area of about 311 square 
kilometers to a depth of 13.5 meters. This Plei- 
stocene lake was named Lake Cochise by O. E. 
Meinzer (1913:34), and then as now, had no 
outlet. An extensive wind-built sand and clay re- 
gion lies south and west of Willcox. This region 
is characterized by scattered ridges and hills, 
some as high as 15 meters and several kilometers 
long, laid out in no specific pattern, but sepa- 
rated from each other by ponds or depressions 
which fill with water during the rainy season 
and are capable of holding this water for a con- 
siderable time thereafter. 

The isolation of the Sulphur Springs Valley, its 
unique drainage system which is partially south 
but mostly internal, the peculiarities of the top- 
ography where water can remain in blowouts 
for extended time are factors which contribute 
to the valley's ability to sustain a large diversified 
tiger bettle fauna. 


SYSTEMATIC ACCOUNT 
Genus Cicindela 
Cicindela willistoni sulfontis Rumpp, new sub- 


species. 
(Figure 1a.) 


DESCRIPTION.—Medium in size, narrow, con- 
vex, robust in form; color dark blue-green, green- 


RUMPP: TIGER BEETLES 


ish bronze, or brown; dull above, brilliant below; 
maculation broad, connected along edge of elytra 
HEAD: A few white hairs on vertex just above 
frons in interocular space. ELYTRA: The macula- 
tion consists of a broad C-shaped humeral lunule. 
The middle band is extended upward along the 
edge of the elytra to meet the humeral lunule, 
also considerably downward to join the apical 
lunule; the band enters the edge with only a 
slightly downward direction, then bends suddenly 
so that it is nearly parallel to the suture for a 
long distance before it ends in an inwardly di- 
rected bulb; the apical lunule is broad at the tip 
with its inner edge extending outwardly and 
down toward the edge where the lunule bulges 
without recurving. DIMENSIONS: Holotype male, 
green - length 13.0 mm, width 4.8 mm. Allotype 
female, brown, similar to the male but broader - 
length 13.0 mm, width 5.4 mm. 

ETYMOLOGY.—This subspecies is named for 
the Sulphur Springs Valley by combining the Latin 
sulfo (sulphur) with fontis (a spring). 

THE TYPE SERIES.—Until 1965 the type series 
consisted of only 4 specimens collected between 
5 and 9 kilometers southeast of Willcox, in a re- 
gion of wind carved sinks. In August of 1969 Mr. 
Jay M. Sheppard discovered the primary habitat 


171 


at which time large series were obtained. The 
type location is 5.6 kilometers west-southwest of 
Willcox, on the playa, and a few hundred meters 
south of the tracks of the Southern Pacific Rail- 
road. All paratypes from the north shore of the 
playa, as all those from the blowouts to the east 
are considered paratypical topotypes; all speci- 
mens are assumed to be from the same general 
population. 

The type series consists of 217 specimens dis- 
tributed as follows: Holotype, male, type no. 
12,530 in the collection of the California Academy 
of Sciences; allotype, female, in the N. L. Rumpp 
Collection (NLRC). There are 215 paratopotypes 
of which 68 are in the NLRC, 86 in the collection 
of Rev. Bernard Rotger, C.R., of Pagosa Springs, 
Colorado, 53 in the collection of Jay M. Shep- 
pard of Laurel, Maryland, and 2 each in the col- 
lections of the American Museum of Natural His- 
tory (AMNH), New York City, the U.S. National 
Museum (USNM), Washington, D.C., and the 
California Academy of Sciences (CAS), San Fran- 
cisco, California. 

VARIABILITY.—The elytral maculation is strik- 
ingly constant, but the color is either green or 
brown with few intermediates. This is shown in 
Table 1. 


TABLE 1. Distribution of color and sexes in the type series of C. willistoni sulfontis. 
green brown intermediate 
ponector Date male female male female male female 
F. H. Parker 13:X:1949 1 
N. L. Rumpp 2:X:1961 1 1 
N. L. Rumpp 13:X:1964 1 
G. Gaumer VII:1969 1 
J. M. Sheppard 10: VIII:1969 2 5 5 3 
B. Rotger 17: VII1:1969 28 Di, 28 11 1 3 
N. L. Rumpp 20:VII:1970 6 12 11 10 
J. M. Sheppard 20:VII:1970 13 14 2. 10 3 
N. L. Rumpp 8:VII1:1971 1 6 1 
Subtotals 51 67 57 34 mA i 
Totals (217 specimens) aon 91 8 


172 


COMPARISON WITH OTHER SUBSPECIES OF 
C. WILLISTONI.—AII specimens of the type series 
are similar in marking and reasonably homogen- 
eous in size, but their color varies. In the majority, 
the color is blue-green, similar to ssp. pseudosen- 
ilis and ssp. praedicta of the Death Valley Hydro- 
graphic System of California and Nevada, but 
unlike these, ssp. su/fontis has a few hairs be- 
tween the eyes, but not nearly so many as are 
found on the more eastern subspecies willistoni, 
hirtifrons and estancia. A lesser number, making 
up nearly the balance of the population, is 
brown or bronze in color, similar to the eastern 
subspecies. The only known location for ssp. 
sulfontis is near Willcox where it is isolated by 
great distances from all other populations of 
C. willistoni. This isolation is especially signifi- 
cant because C. willistoni is usually not found in 
the Colorado River drainage system! as reported 
by Rumpp (1961: 168), although it has been 
located now in Arizona, a fact not known at that 
time. Cicindela willistoni shows a preference for 
lacustrine salt flats, although it may also invade 
marshy areas and semi-dry river beds where 
saline muds are exposed. Depending on eleva- 
tion and temperature, it may emerge early in the 
year at lower elevations, much later at higher 
altitudes. In the Sulphur Springs Valley the cli- 
mate is normally dry in May and June and re- 
mains that way until the start of the rainy sea- 
son in mid-July, at which time ssp. sulfontis sud- 
denly appears in numbers on the Willcox Playa 
where it remains in evidence throughout the 
rainy season. The larva builds a turret on the 
playa in the fashion of ssp. hirtifrons as reported 
by Willis (1967:176-177). 


DISTRIBUTION OF CICINDELA WILLISTONI. 
—Cicindela willistoni has an extensive distribu- 
1ion in western United States. The species ranges 
throughout the Great Basin and that part of the 
Basin and Range province extending into south- 
eastern Arizona, but nearly exclusive of the Colo- 
rado River drainage system. East of the Rocky 
Mountains it occurs in Wyoming, Kansas, Okla- 
homa, Texas, and New Mexico. It may be found 
wherever there are wet playas, defined as sea- 
sonal salt flats or perennial lake beds. The various 
subspecies from the eastern side of the distribu- 
tion are a bright bronze color and bear a deeper 


1An exception exists in the Ft. Bridger, Wyoming popula- 
tion of ssp. willistoni. This area is connected hydrographically 
to the Green River, the main tributary of the Colorado River. 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 4 


middle band, usually with very broad to conflu- 
ent maculation. In these the head always bears 
a number of long decumbent hairs (Willis, 1967: 
303). They may be geographically distinguished 
as ssp. willistoni of Wyoming; ssp. hirtifrons of 
western Kansas, Oklahoma, Texas, and north- 
eastern New Mexico; ssp. estancia the endemic 
population of the Estancia Valley in central New 
Mexico, and finally ssp. funaroi of Jemez Creek 
in northcentral New Mexico. In the Great Basin, 
most populations are of ssp. echo. These may be 
subdivided into the eastern or Bonneville echo 
populations, and the western or Lahontan am- 
adeensis populations, with such little differences 
in habitus and color as to be considered synony- 
mous (Cazier, 1936:157). The name_ spaldingi 
was proposed by T. L. Casey (1924:14) for one 
bright green, broadly maculated specimen from 
Calleo, Utah. This is in a region encompassing 
widely variable populations of ssp. echo. These 
populations were sampled by the author on 29 
May 1970 at Blue Lakes, Tooele County, Utah, 
at the edge of the Great Salt Lake Desert, ap- 
proximately 80 kilometers northwest of Calleo. 
Another location is at Fish Springs Wildlife 
Refuge, Juab County, Utah, also at the edge of 
the Great Salt Lake Desert and 25 kilometers 
east of Calleo. This last site was visited by the 
author on 4 May 1962, and by Lawton and Willis 
(1974:51) on 24 July 1971. These populations 
are extremely variable in that a few individuals 
are like Casey’s spaldingi, while others resemble 
ssp. estancia; however, the great majority of 
specimens is typical of ssp. echo. The third larg- 
est hydrographic entity of the Great Basin is the 
Death Valley system where C. willistoni is inter- 
spersed with populations of ssp. echo, ssp. 
pseudosenilis, and ssp. praedicta. Here colors 
trend to green or dark blue-green, with or with- 
out maculation. In all of the Great Basin popula- 
tions from Utah and Nevada, southern Idaho, 
and eastern parts of Oregon and California, the 
vertex bears only a few hairs or none at all. 


Cicindela pulchra dorothea Rumpp, new sub- 
species. 
(Figures 1b, 2.) 


DESCRIPTION.—Large in size, form robust; 
head, prothorax and elytra brilliant metallic-cerise 
or wine-red. Elytra with definite maculation, some- 
times fully connected along sides. HEAD: Brilliant 
red on vertex from eye to eye, fringed with gold 
and green before shading into the bright deep 


RUMPP: TIGER BEETLES 


73 


[On 


Figure 1. Dorsal views of: a. Cicindela willistoni sulfontis; b. C. pulchra dorothea; and c. C. nevadica citata. 


Scale indicates 10 mm. 


blue that covers the underside. Slightly rugose on 
the vertex, wrinkled near the eyes; deep setiger- 
ous punctures on the frons. Vertex with sparse 
erect white hairs, frons covered with long white 
decumbent hairs, genae bare. First four segments 
of the antennae bright green; first segment with 
over twenty stiff hairs on the outer edges, sec- 
ond segment bare, third and fourth with a 
sparse row of white hairs on the outer edges, 
with one setigerous puncture on the inner tips. 
Clypeus bright blue-green, with two setigerous 
punctures. Labrum white with a narrow dark 
edge, tridentate, with six setigerous punctures 
bearing long white hairs close to the outer edge. 
THORAX: Pronotum as broad as the head ex- 
cluding the eyes, broader than long by a ratio 
of 4 to 3; broadest at the forward one third, 
then slightly tapering inward along a slight 
curve to the rounded basal angle. Surface finely 
rugose, brilliant red fringed with gold and green. 
Anterior and basal impressions deep, bright blue 
fore and aft respectively of these impressions; 
median impression very narrow, shallow, colored 
green. ELYTRA: Brilliant metallic red throughout, 
except for the very narrow outer edge which is 
blue with a row of green punctures. Surface 
punctured, these punctures deepest nearest the 


base, very shallow toward the apex. Suture 
golden-green. Maculation complete but reduced 
in extent; central band triangular, broadest at 
the edge of the elytra, very slightly recurved 
downward, reaching nearly half way to the su- 
ture; humeral lunule shallow at its tip, some- 
times connected to the central band; apical lun- 
ule narrow, ending in a shallow dot, then nar- 
rowly connected to the middle band. UNDER- 
SIDE: Bright deep blue. Proepisterna covered 
with long decumbent hairs; meso- and metaepis- 
terna with shorter decumbent hairs. Upper ab- 
dominal segments with sparse white hairs on the 
sides, and long white hairs at the bottom edge 
of the 3rd, 4th, and 5th sternites; pro- and meso- 
coxae with long recumbent hairs, metacoxae 
bare; trochanters bare except for one setigerous 
puncture at the tip of those of the front and 
middle legs; femora blue with long hairs, more 
so on the anterior ones than on the middle and 
back ones; tibiae blue-green with rows of stiff 
white bristles, apical spurs about two thirds the 
length of the first tarsal joints; tarsal joints with 
several erect hairs, especially at tips; tarsal claws 
unpigmented, short, about half the length of the 
fifth tarsal joints. DIMENSIONS: Male - length 15.2 
mm, width 6.0 mm. Female - length 15.6 mm, 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 4 


Figure 2. Habitat of Cicindela pulchra dorothea south of Williston, Arizona on 26 July 1964. In the distance is a 


water filled pond. 


width 6.4 mm. 

ETYMOLOGY.—This insect is named in mem- 
ory of my late wife, Dorothy, who collected many 
of the specimens in the initial large series. 

THE TYPE SERIES.—There are 78 specimens in 
the type series. This includes the holotype male, 
the allotype female, and 44 topotypical paratypes 
collected by N. L. and D. H. Rumpp at the type 
location 5.2 to 6.2 kilometers southeast of Will- 
cox, on 26 July 1964, and an additional specimen 
collected on 11 September 1965. Five paratopo- 
types were captured on 21 May 1970, three on 16 
July 1974, twelve on 17 July 1974, and one more 
on 6 August 1975, all by the author. Two para- 
topotypes were collected by Dr. John Stamatov 
on 8 August 1971. The series also includes one 
paratype from 11.3 kilometers west of Douglas, 
Arizona, dated 6 August 1965. 

Distribution of the type series: Holotype male, 
type no. 12529 in the California Academy of Sci- 
ences; allotype female, 66 paratopotypes and 
one paratype in the NLRC. Two paratopotypes 
deposited in each of the collections of the 
AMNH, the USNM and the CAS. Two are in the 
private collection of Dr. John Stamatov. 

COMPARISON WITH C/ICINDELA PULCHRA 
PULCHRA.—From the time it was described by 
Thomas Say in 1823, Cicindela pulchra was con- 


sidered a monotypic species from midwestern 
United States, ranging from Montana to Texas. 
Subspecies pulchra is larger, generally immacu- 
late, with a color varying from dark red to deep 
violet. Some specimens, and these are few, have a 
triangular marginal middle spot not extended at 
the sides, sometimes with two dots representing 
a vestigial humeral lunule; otherwise the border 
of the elytra is broadly blue. Subspecies dorothea 
has a brighter red color, often with strong green 
reflections when viewed from an angle. In many 
specimens the white maculation is connected 
along the outer edge of the elytra, while in a 
few the maculation is reduced to dots. Less than 
ten percent of the specimens in the type series 
have a narrow middle band with blue connec- 
tions along the outer edge of the elytra, and 
whereas these resemble the rarely maculated 
specimens of ssp. pulchra, the color is typical of 
ssp. dorothea, as is the smaller size. 

Although ssp. dorothea is now isolated in 
southeastern Arizona, it was probably derived 
from the ancestral Rio Grande system during the 
Mio-Pliocene when the flow of the ancestral 
upper Colorado River drained east, and the Con- 
tinental Divide was west of the valley, as will be 
discussed later. Now that this subspecies occu- 
pies a region that drains into the Yaqui River 


RUMPP: TIGER BEETLES 


system, it will probably be found in northern 
Sonora, especially since it has been located a few 
meters from the Mexican border west of Douglas, 
Arizona. 

Some New Mexico specimens of C. pulchra 
from the USNM, AMNH, and private collections 
were examined. Those from the northwest or 
Farmington district can be identified with ssp. 
pulchra, whereas those from the southwest or 
Silver City district appear to identify with ssp. 
dorothea, as do those from the trans-Pecos re- 
gion of western Texas. 


Cicindela nevadica citata Rumpp, new subspecies 
(Figure 1c.) 


DESCRIPTION.—Medium size, body narrow, 
color brownish bronze with green micro pits, 
maculation of elytra narrowly connected along the 
outer edge. HEAD: Small relative to body size but 
with salient eyes; finely granulate vertex, frons 
very narrowly striate. Color green suffused with 
bronze. Pilosity of medium density on vertex, 
frons, and clypeus. First 4 joints of antennae 
green with bronze reflections on last 3 joints; 
first joint with 12 to 15 white recumbent hairs, 
2nd joint bare, 3rd and 4th with 2 to 4 short 
hairs on outer edges and 2 setigerous punctures 
at tips. Clypeus white, barely produced in front, 
unidentate, ratio of width to length is 3 to 1; it 
bears 16 to 18 setigerous punctures located near 
anterior edge in two irregular rows. THORAX: 
Pronotum narrow, about as broad as the head 
excluding the eyes, cylindrical, length equal to 
width, finely rugulose, color greenish bronze. 
Anterior impressions of medium depth, basal 
impressions deeper, median impression shallow; 
sparse white decumbent hairs cover the top sur- 
face, more so at the edges, top, and bottom leav- 
ing the disc nearly bare. ELYTRA: Long, narrow, 
widest at the middle and twice as wide as the 
pronotum. Male - sinuous near tip, finely serrul- 
ate in this region; tips end in spines. Female - 
more sinuous and indented; terminal spines are 
smaller. Color brown except for center half from 
suture Outward which is bronze; pitted through- 
out, pits green. A row of small green punctures 
descends near the suture, two rows of small 
punctures are located in the humeral impres- 
sions. The maculation is white and of typical 
nevadica design, except that the middle band is 
more narrow where it enters from the outer 
edge, and the central extension is longer; all 
markings are narrowly connected along the sides. 


WHS 


UNDERSIDE: Mostly shiny brown with green re- 
flections, however the genae are bright green, 
narrowly striate and covered with dense decum- 
bent white hairs. The proepisterna are bronze 
with green reflections, densely covered with 
long white decumbent hairs; meso- and metaepi- 
sterna similarly clothed with long white hairs; 
pro- and mesocoxae also clothed with long white 
hairs, metacoxae bare; trochanters unpigmented, 
bare except for a setigerous puncture at tip of 
pro- and mesotrochanters. All abdominal sternites 
clothed with decumbent hairs on outer halves, 
with a few hairs at forward edges centrally. Fore 
and middle humeri with several rows of dense 
white hairs on the outer and inner sides; pos- 
terior humeri sparsely covered with non-over- 
lapping hairs, none on outer sides. Tibiae bronze 
with greenish reflections, sparsely bristled. DI- 
MENSIONS: Male - length 10.3 mm, width 4.2 
mm. Female - length 11.0 mm, width 4.4 mm. 


ETYMOLOGY.—The name citata alludes to the 
insect’s fast running over moist sandy lacustrine 
shores. 


THE TYPE SERIES.—The type series consists of 
48 specimens. The holotype male was collected 
at the edge of the central playa 8 kilometers west- 
southwest of Willcox on 20 July 1970. The allo- 
type female is from 2.9 kilometers south of Will- 
cox, captured on 8 August 1955. Forty-six para- 
topotypes were collected near Willcox at sites 
such as Twin Lakes, nearby ponds, or at the 
northern edge of the Willcox Playa; all were col- 
lected from July through August between the 
years 1955 and 1975. 

The holotype is deposited in the collection 
of the California Academy of Sciences as type 
no. 12,528. The allotype and 29 paratopotypes 
are in the NLRC. Paratopotypes in private col- 
lections are: Sheppard 3, Clifford 6, and Pearson 
2. Two each were deposited in the AMNH, the 
USNM, and the CAS collections. 


VARIABILITY: Most of the specimens in the type 
series are similar to the described type except for 
two specimens. One is a large female 12.4 mm 
long, with more cupreous coloration, the other a 
very small male only 9.1 mm long. 


COMPARISON WITH OTHER SUBSPECIES OF 
CICINDELA NEVADICA.—The nominate subspe- 
cies nevadica of the Great Basin region of Nevada 
and California is of a coppery bronze with green 
reflections, somewhat similar in color to. ssp. 
citata. The typical form of ssp. nevadica is reputed 
to be from Ash Meadows, Nye County, Nevada, 


176 


although LeConte (1875:159) merely reported 
it from Nevada. In the Nevada populations of 
ssp. nevadica the middle band seldom has any 
extension along the border, whereas in California 
populations there is usually a narrow extension 
along the border, sometimes connecting all the 
markings. The elytral pattern of the California 
populations more nearly matches the woodcut 
accompanying the description by LeConte, but 
he also states in the notes following his descrip- 
tion that the markings were not connected along 
the margin. The markings of ssp. nevadica, 
where there is a connection along the outer 
border, differ from ssp. citata in that the middle 
band, where it enters from the edge, is broader 
in the former than in the latter; furthermore, the 
extension of this band is much deeper and nar- 
rower in ssp. citata. Subspecies tubensis differs 
from ssp. citata in that it has a red color, and al- 
though its maculation is always connected along 
the margin, these markings are much broader. 
Subspecies knausi of the upper Rio Grande basin 
is more nearly like ssp. citata in markings, but 
it also has slightly broader markings and _ less 
greenish coloration. Subspecies citata fits best 
between ssp. nevadica in color, and ssp. knausi 
in markings. The range of ssp. citata is the Sul- 
phur Springs Valley, possibly into the Yaqui 
River system of Sonora. Those from the north- 
ern shores of the Gulf of California, specifically 
from Puerto Penasco, can be identified with this 
subspecies. 


ECOLOGICAL ASSOCIATIONS OF CICINDELA 
SPECIES OF THE SULPHUR SPRINGS VALLEY 


There are seventeen species of the genus Ci- 
cindela in the Sulphur Springs Valley. These are 
separated below according to their ecological as- 
sociations. Comparisons of related species and 
subspecies are given for some of the seventeen. 


On wet flats, at the edge of water 


1. C. praetextata erronea. This bright green or 
blue-green subspecies is endemic to the valley. 
It may be found on the northern shore of the 
Willcox Playa, and from the wind-formed ponds 
to Croton Springs. It is strikingly different in 
color from the bronze ssp. praetextata of the 
Colorado River drainage system, and from the 
red ssp. fulgoris of the Rio Grande system and 
upper regions of the Little Colorado River. 

2. C. marutha marutha. When individuals of 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 4 


this subspecies emerge at the start of the sum- 
mer rains, they form the largest populations along 
the mud flats of the ponds near Willcox. They 
will also invade marshy grasslands for they are 
not restricted to open areas, and at night they 
will come to light. This population is made up 
of nearly all pure grass-green individuals. Only 
4 percent of the total are red. Most other pop- 
ulations of C. marutha in Arizona and New Mex- 
ico contain a high percentage of red individuals, 
but the northern ones contain the highest per- 
centage of this color. In southeastern Utah there 
are pure red populations which should be re- 
ferred to ssp. rubicunda. 


3. C. haemorrhagica. The valley’s population 
contains rather small, well marked individuals 
which have most often been identified with ssp. 
woodgatei. There is a resemblance to C. cartha- 
gena carthagena, especially in the median widen- 
ing of the elytra and the angle so produced 
where the outer edges converge toward the tip. 
Individuals are found at the edge of water, usu- 
ally on the darker spots where they segregate 
from other species. 


4. C. nevadica citata. Individuals are relatively 
scarce in the valley, most being found immedi- 
ately south of Willcox. This scarcity may be at- 
tributed to the high salinity of the local flats, be- 
cause elsewhere this species seems to prefer 
damp sand, free from salt incrustations. 


In grassy marshlands and very wet spots 


5. C. tenuisignata. This is a brown monotypic 
species common to the Rio Grande system, the 
lower Colorado River drainage system, and Son- 
ora. Individuals prefer damp as well as wet sandy 
soil. 

6. C. punctulata chihuahuae. This is a grassland 
tiger beetle that is relatively rare in the valley. 
When found it will be in wet places. Individuals 
exhibit a variety of colors, from a bronzy green 
to a bright blue, with or without maculation. 
Bluish green is the most common color. 

7. C. nigrocoerulea nigrocoerulea. This sub- 
species is polymorphic with individual members 
exhibiting a variety of colors that range from dull 
green to deep blue to black; blue-black is the 
predominant color. Individuals may be found 
most frequently in the thicker marshy areas and 
muddy spots, sometimes in great abundance. 

8. C. horni horni. This is also a polymorphic 
subspecies with colors varying from metallic 
green or bright blue, to black. It is widely dis- 


RUMPP: TIGER BEETLES 


tributed in the valley from Douglas to Willcox, 
from the lowest grassy areas to upland meadows. 

9. C. sedecimpunctata sedecimpunctata. This 
is a very common subspecies in southern Ari- 
zona, Sonora, and the Rio Grande system. It has 
a tolerance for high altitude, having been located 
in the Pinalino Mountains at elevations of 2700 
meters, but always near ample water supplies. 
Although it is related to, and resembles C. 
haemorrhagica, individuals are noticeably smaller 
than in that species. 


On muddy flats of the Willcox Playa 


10. C. willistoni sulfontis. Members of this 
subspecies emerge in great numbers on the 
playa south-southwest of Willcox after the rains 
begin in mid-July and the surface of the playa 
becomes mucky. This form was first discovered 
at one of the ponds in the wind-built area south 
of Willcox, an area which it seldom invades. 
One individual was captured here following a 
long flight from the edge of a pond over a dry 
clay spot populated by C. pimeriana. Individuals 
are strong fliers, making frequent flights of as 
long as 10 meters, with very brief rests between 
flights. 


On dry flats, not far from water 


11. C. pimeriana. This monotypic species is the 
most common one found in early spring and 
again in autumn. Its characteristic color is brilliant 
metallic green, although the few spring individ- 
uals that survive until October become dull black. 
This tiger beetle is found only in the Sulphur 
Springs Valley and the San Bernardino Valley to 
the southeast. This last area contains the type 
locality at the San Bernardino Ranch (the old 
Slaughter Ranch) on the Mexican border. The 
species is related to C. pulchra, with which it 
shares a general area south of Willcox, but its 
preferred habitat is on bare clay banks near 
water. An individual’s flight is long and singular, 
and it makes such flights between long rest 
periods, possibly as a defense against asilid rob- 
ber flies that prey upon it. The fly’s attack strategy 
is to swoop upon the beetle as the beetle takes to 
flight. The fly then embraces its prey with its legs 
while still flying and inserts its beak at the junc- 
ture of the open elytra, directly into the meso- 
sternum, stunning the beetle almost immediately. 
The flight of the asilid is so slowed by this load 
that fly and prey can be netted with ease. Five 
captures were made with C. pimeriana as the fly’s 


WAZ 


victim, and once with a C. marutha as victim. 
12. C. ocellata ocellata. This small dark-brown 
subspecies belongs to the Sonoran fauna and is 
widely dispersed throughout southern Arizona. It 
differs most noticeably from ssp. rectilatera of the 
Chihuahuan fauna by its rufous abdomen. 


In grasslands, away from water 


13. C. pulchra dorothea. This strikingly beauti- 
ful tiger beetle is found exclusively on open flat- 
lands covered with clumps of grass. The beetle 
emerges from these clumps to fly onto bare 
spots in one long flight. There it will either re- 
main in the open where it will seek the shade of 
grass, or it will rapidly run into the nearest clump 
of grass and disappear from view. 

14. C. lemniscata lemniscata. This small red 
tiger beetle with pale legs and longitudinal macu- 
lation seeks open spots far from water. When dis- 
turbed it quickly takes to wing, but it is a weak 
flier with an erratic flight. Although often seen in 
the daytime, its habits are more nocturnal than 
diurnal, and at night it will come to light in great 
numbers. 

15. C. debilis. This small dull-green tiger beetle 
is related to C. /Jemniscata, but unlike that species 
it rarely flies. It prefers heavy grass where it 
seeks the densest parts for refuge. It is rarely en- 
countered in the daytime and may be nocturnal 
in habit. The species will usually not fly, but 
when coaxed to do so its flight is erratic and very 
short, only a few decimeters in distance. The 
green C. debilis segnis was described as a variety 
from southern Arizona by Harris (1913:69), but 
Cazier (1954:287), after studying large samples 
from southern Arizona and Durango, Mexico did 
not favor retaining this subspecies. A population 
of darker individuals of this species was found 
near Willcox, and one with brighter green indiv- 
iduals was found near Douglas. However, these 
color variations may have been due to the age of 
the individuals sampled, the older possibly being 
the darker. 

16. C. obsoleta santaclarae. This is the largest 
tiger beetle in the valley, often measuring 17 mm 
in length. It too is polymorphic for it may be 
green, blue, brown, black or red, although the 
predominant color seems to be green. The elytra 
are spotted, and a vestigial middle band is in- 
dicated by two dots on each elytron. The species 
is robust and its members are excellent fliers. Its 
flight is long, sometimes on the order of 100 
meters, and often very high. It is found in grass- 
lands on well-drained alluvial slopes, particularly 


178 


along the edges of the valley. In the grasslands 
west of Douglas it is encountered with C. nigro- 
coerulea, C. pulchra, C. horni, C. lemniscata, and 
C. debilis. 


Tree areas near water 


17. C. viridisticta arizonensis. This is another 
very small tiger beetle related to C. debilis and 
C. lemniscata. Near Douglas it is found among 
grasses on the slopes of drainage ditches in the 
chaparral-covered plains, while at Kansas Settle- 
ment it is found in grass near willows and tules 
that grow at the sides of irrigation canals. The 
species is a weak flier with a short and erratic 
flight. It prefers to remain on the ground where 
it will move with great speed and agility through 
the grass. 


KEY TO THE SPECIES OF CICINDELA 
IN THE SULPHUR SPRINGS VALLEY 


Tas Abdominall Segments MUfOUS jn. cccnccesncec-scneqeececreenreneneceec- 2 
1b. Abdominal segments black, dark blue or greenish .... 4 


2a. Elytra gradually widening to apical fourth; color 
[eA] cecsceRerceckeunbe Pe GeS-er acaeeccaen ce USO Are rae asc te paneer cocECeREMURESE aS 3 

2b. Elytra widest at middle or basal third; color black 

haemorrhagica 


3a. Post, median, lateral and apical dots present ............ 
2 ID PRIOR RE Ren ee ee ae ere sedecimpunctata 
3b. Post, median, lateral and apical dots absent ....ocellata 


4a. Front trochanters only with seta (small species, less 
aetna, G3 Inenn) (ROG) coetee cto coeceoscecet tebe ccaceeeeeren eran ceeeeEceoceeed nec 5 
4b. Front and middle trochanters with seta .................... 6 


5a. Elytra bright red, shiny, deeply punctate; markings 
consisting of a longitudinal line .................... lemniscata 

5b. Elytra dull grayish brown with a longitudinal row of 
green foveae; middle band and apical lunules present 
Blea eater) PEE EES Se eee eae eee rrere ren ere viridisticta 


6a. Small species, less than 9 mm long (velvety green 
in color, elytral maculation consisting of a wide 


partially submarginal border) ................--.-...------ debilis 
6b. Larger species, 10 mm long or longer ................--..--.- 7 
7a. Genae covered with decumbent hairs .................2....-.-- 8 
IMMA Clic POR ARG Mesto ittees cee ccaceseec eee ceree scesuere wcrenneecnsesteccenersct nae 9 


8a. Antennal scape with subapical sensory hairs only; 
bright green, occasionally red (elytra with feeble 
SVanieelt Gpollats)) aecceescocoreececucecceasoreo-c- pees eneeceercReencec ance marutha 

8b. Antennal scape with decumbent hairs below the sub- 
apical setae; brown or bronze with greenish reflec- 


HONS ons cc cecec cs fara sone cee ter aseene errno eee etecenedesesecteencs nevadica 
9a. Frons or vertex to some extent pilose ..........0....0..00--- 10 
9b. Frons and vertex bare except for ocular setae ............ 12 


10a. Pilosity of vertex sparse, frons bare; color dark 
green or brown; (maculation complete and joined 


Ate MVANBIN) | sssecesees-aceecece ceases cecsevonteccontorenecscteeeteeres willistoni 
10b. Pilosity of frons and vertex medium to dense; color 
yrs ren ie ees ae noe re Ae ee 11 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 4 


41a. Green or blue-green; immaculate or with very small 
apical dots, rarely any median dots ................ pimeriana 

11b. Red; maculation marginal consisting of humeral, 
median, and apical spots, sometimes connected; 
(Keren Si Ze) ett cs cvcessceseese reese stcecseceeceeotese meneeeneee pulchra 


12a. Under surface sparsely to densely pilose laterally......13 
12b. Under surface bare except for a few scattered hairs, 
especially on sides of metasternum ...............-.....+ horni 


13a. Proepisterna densely clothed with long white decum- 
bent hairs; maculation complete .................. praetextata 
13b. Proepisterna bare or sparsely clothed with hairs ...... 14 


14a. Elytra with subsutural row of blue foveae from base 


fO!) AD OX cece. ccnceccedncnc-caeceenss-cessnesse<cvacenna beaneteseeaee ee AS 
14b. Elytra with shallow or obscured foveae, or none at 
ANE occ cede c Sees sdcadadou seeds sates SoeSSy cece eee oe eee 16 


15a. Color green or blue-green; elytra narrow and slightly 
widening to apical fourth, microserrulate near apex; 
maculation consisting of small dots except for a nar- 
TOWaaPiGal WM ee eeee sea c cece ean eee eee aeeneee punctulata 

15b. Color dull green, dark blue or blackish blue; form 
robust; elytra evenly arcuate, broadest at middle, not 
serrulate near apex; maculation usually absent ........ 
BREE PR SEC een SEER Se Ree a SEA ee bisects ance nigrocoerulea 


16a. Elytra with shallow or obscured foveae; maculation 
narrow and complete, descending portion of middle 


band) long-msimalllenminieSizemeseseeneteneeesereee tenuisignata 
16b. Elytra without foveae; maculation consisting of inter- 
ruptediidots* Very) langel lin SIZe) ster eccpcentseeneeeee obsoleta 


CLASSIFICATION OF THE SULPHUR SPRINGS 
VALLEY CICINDELA 


By using the characters contained within the 
male genitalia in conjunction with a few external 
characters, Rivalier (1954) arrived at a natural 
classification of the genus Cicindela. Based partly 
on his system, the species of the Sulphur Springs 
Valley are tentatively classified below. Some col- 
lecting locations in the Sulphur Springs Valley are 
indicated in brackets for each species. 


Subgenus Cicindela?” 


C. pulchra group 
C. pimeriana LeConte [Willcox, Douglas, Croton Springs] 
C. pulchra dorothea Rumpp [Willcox, Douglas] 


C. willistoni group 
C. willistoni sulfontis Rumpp [Willcox] 


C. punctulata group 
C. tenuisignata LeConte [Willcox] 
C. punctulata chihuahuae Bates [Willcox] 


C. rufiventris group 
C. haemorrhagica woodgatei Casey [Willcox, Kansas Settle- 
ment] 
C. sedecimpunctata sedecimpunctata Klug (Willcox, Doug- 
las] 
C. ocellata ocellata Klug [Willcox, Douglas] 


2Includes all of Rivalier’s (1954) Cicindela and Cicindelidia. 


RUMPP: TIGER BEETLES 


C. obsoleta group 
C. nigrocoerulea nigrocoerulea LeConte (Willcox, Douglas, 
mountain slopes] 
C. obsoleta santaclarae Bates [mountain slopes, Douglas] 
C. horni horni Schaupp [Willcox, Elfrida, Douglas, moun- 
tain slopes] 


Subgenus Habroscelimorpha 


C. circumpicta group 
C. praetextata erronea Vaurie [Willcox, Croton Springs] 


Subgenus Cylindera® 


C. nevadica group 
C. nevadica citata Rumpp [Willcox] 
C. marutha marutha Dow [Willcox, Croton Springs] 


C. lemniscata group 
C. lemniscata lemniscata LeConte [Willcox, Dos Cabezas, 
Douglas] 


C. celeripes group 
C. viridisticta arizonensis Bates [Willcox, Kansas Settlement, 
Douglas] 
C. debilis Bates (Willcox, Douglas, mountain slopes] 


DERIVATION OF THE C/CINDELA 
IN THE SULPHUR SPRINGS VALLEY 


GENERAL. —An underlying assumption made 
in this section is that tiger beetles of the arid re- 
gions of the Southwest were dispersed by way of 
hydrographic paths connecting various basins dur- 
ing wetter periods, or so-called pluvial times. 
These periods of higher rainfall were generated 
during glacial cycles when storm cyclones moved 
south ahead of advancing ice sheets in the north. 
Another assumption is that orogeny was also a 
force that had a direct impact on the direction of 
rivers and the distribution of rainfall. From the 
present distribution of the Cicindelidae in the 
Southwest, coupled with some data on past geo- 
morphology, a number of speculative routes can 
be inferred. 


EASTERN COLORADO-RIO GRANDE HYDRO- 
GRAPHIC CONNECTIONS. —Most of the species 
in the valley were derived from the Rio Grande— 
Chihuahuan fauna. To better understand this 
strong influence from the east, it is essential to 
realize that the Colorado River drainage basin of 
today did not always drain into the Gulf of Cali- 
fornia. There is much evidence indicating that 
during mid-Cenozoic (Miocene) the drainage pat- 
terns were to the east. The latest review of the 
evolution of the Colorado River explains this pat- 
tern. The following partial quotation is from 
McKee et al. (1967: 54): 


3Includes all members of the subgenus Ellipsoptera Dokh- 
tourow. 


179 


“General drainage patterns in northern Arizona during 
Stage 3 (mid-Cenozoic drainage patterns) — At least two 
different drainage pattern systems, separated by the 
Kaibab upwarp, are believed to be present. The eastern 
system, here called the ancestral upper Colorado system, 
in inferred to have flowed southward from Utah then 
southeastward across northeastern Arizona approximately 
along, or somewhat north of, the present Little Colorado 
River course, but in a reverse direction. . . . This drain- 
age may possibly have connected with an ancestral Rio 
Grande drainage flowing to the Gulf of Mexico.”’ 


The distribution of Cicindela along the upper 
course of the Little Colorado River prompted the 
following statement by Rumpp (1961: 180): “The 
Little Colorado River was created during, and as 
a result of uplift. Prior to this, its upper basin was 
part of an eastern hydrographic system.” This 
conclusion, although not as encompassing as that 
of McKee et al. is nevertheless similar. It was 
based on the occurrence of several species of 
Cicindela along the upper Little Colorado River, 
such as the large red C. hirticollis (ssp. ponder- 
osa), and also C. sperata, C. nigrocoerulea, C. ful- 
gida and C. /epida, all of which occur in the Rio 
Grande system. C. fulgida and C. lepida have 
spread as far as Utah. In the Rio Grande system 
C. fulgida displays a longer descending middle 
band, a characteristic that is also common to 
those populations of the upper Little Colorado 
River basin. 

With the ancestral upper Colorado River drain- 
ing eastward from mid-Miocene to some later 
undetermined time, it follows that the Contin- 
ental Divide was farther to the west than it is 
today. This may have been the situation from 
Montana to southern Arizona. In this latter region 
the divide would have been west of the Sulphur 
Springs Valley such that the regional drainage 
would have been eastward eventually reaching 
the Gulf of Mexico, possibly as a port of the an- 
cestral Rio Grande River system. Along such a 
hydrographic pathway, all of the middle Rio 
Grande species now found in southern Arizona 
could have migrated westward. These include 
C. pulchra, C. tenuisignata, C. punctulata, C. 
nigrocoerulea, C. horni, C. obsoleta, C. marutha, 
C. willistoni, C. sedecimpunctata, C. ocellata, C. 
nevadica, C. lemniscata. and C. praetextata. One 
species, C. haemorrhagica, is of a more Sonoran 
type which may have migrated eastward at that 
time. During this early migration period the 
species probably met few new conditions, but 
many environmental changes came in time. Be- 
cause of varying ecologies the species adapted 
differently. Some subspeciated along the length 
of the migratory path, while others did so only at 


180 


the extremes of the range. For example, C. pul- 
chra dorothea subspeciated along the whole 
length of its southwestern expansion. Now that 
its range has been disrupted it remains as the 
same subspecies in the Sulphur Springs Valley, 
southern New Mexico (Silver City region), and 
trans-Pecos Texas (Alpine—Marfa region). The 
same holds for C. nigrocoerulea, C. horni, C. sed- 
ecimpunctata, and C. punctulata. Others that sub- 
speciated only at the ends of this long route are 
C. ocellata, C. obsoleta, C. lemniscata, and C. 
praetextata. The C. ocellata representative of the 
middle Rio Grande River region is ssp. rectilatera, 
While in the same region C. obsoleta is repre- 
sented by ssp. vulturina, both significantly differ- 
ent from their western counterparts. Cicindela 
lemniscata also has subspecies in the eastern and 
in the western parts of the range, both subspecies 
having extended their ranges sufficiently to have 
created a hybrid zone along the middle Rio 
Grande River region. Cicindela praetextata ful- 
goris is usually red but sometimes greenish with 
broad markings, while the common western ssp. 
praetextata is brown or bronze with narrower 
markings. The isolation of ssp. erronea next to a 
saline lake bed was described by Rumpp (1957: 
149) as an exceptional case for C. praetextata; 
this isolation possibly explains the startling con- 
trast in color between this oright green sub- 
species and the other subspecies. Cicindela tenu- 
isignata is the only species that appears not to 
have diverged locally, retaining its monotypic 
form throughout its extensive range. 


WESTERN COLORADO RIVER CONNECTIONS. 
—When the upper Colorado River flowed east 
(middle to late Miocene and possibly well into 
the Pliocene) there were normal north-south 
mountains in the region now occupied by the 
Grand Canyon and southwestern Utah. The Hur- 
ricane Fault marks the location of one such 
mountain range. This is specifically mentioned 
here because it is along this ancient fault that the 
large apterous and nocturnal tiger beetle Ambly- 
cheila schwarzi can be found, one population at 
the southern end of the fault at Peach Springs, 
Arizona (type locality), the other at the northern 
end in Diamond Valley, some 20 kilometers north 
of St. George, Utah. Between these populations 
flows the Colorado River through the impassable 
Grand Canyon. It is clear that at one time the 
river was not there, so the population was once 
continuous. If a river existed there in that period, 
it flowed westward. Neither did the Gulf of Cali- 
fornia exist at that time because the Lower Cali- 


PROC. CALIF. ACAD. SCI., 4TH SER., VOL. 41, NO. 4 


fornia peninsular plate had not yet begun its 
movement outward and northward (movement 
did not occur until Pliocene time, about four mil- 
lion years ago). There is strong evidence that 
there was an outlet from this region to the 
Pacific Ocean during the Miocene and possibly 
later. At that time it is probable that the Contin- 
ental Divide precluded the Rio Grande tiger 
beetles from reaching that region. But in time the 
Lower California plate, by its northward pressure, 
cut off drainage to the Pacific Ocean by raising 
the Tehachapi Mountains at the southern end of 
the Sierra Nevada range, thereby creating a re- 
gion of internal drainage. A lake region devel- 
oped during glacial times, eventually ending 
wholly or in part when the Colorado River cut 
through the Grand Canyon and supplied enough 
water to create a discharge to the Gulf of Cali- 
fornia. Trapped populations, originally derived 
from the Rio Grande basin could now expand 
westward wherever conditions would permit. 
Conversely, some western species could now 
migrate eastward into the expanding lower Col- 
orado River basin. 

Westwardly expanding species include C. prae- 
textata praetextata, which in time reached the 
Virgin River Valley as did C. punctulata chihua- 
huae, this latter having been reported also with 
C. tenuisignata at Ash Meadow in the Amargosa 
Desert of Nevada by W. Knaus (1922: 194). 
Whether C. nevadica and C. willistoni ever made 
contact in this manner with their counterparts in 
the Sulphur Springs Valley is unknown; but at 
some time it could have happened, and that pos- 
sibility cannot be discounted. 


YAQUI RIVER HYDROGRAPHIC CONNEC- 
TIONS. —There are two typically Sonoran species 
in the valley. These are C. debilis and C. viridist- 
icta arizonensis. They are not differentiated from 
their Mexican counterparts. Hubbs and Miller 
(1958: 115) found no evidence connecting Lake 
Cochise to the Yaqui River in pluvial times so 
that the present connection of the southern third 
of the valley to the Yaqui River must be of recent 
origin. This leads to the conclusion that these two 
species are recent intruders. 


SPECIFIC ENDEMISM. —The only endemic spe- 
cies in the Sulphur Springs Valley is C. pimeriana, 
a member of the ‘pulchra’ group, and a relative 
of C. fulgida from which it differs in color, lack 
of maculation, sparser pilosity, and a labrum 
which is less produced in front. These differ- 
ences might be viewed as minor but for the fact 


RUMPP: TIGER BEETLES 


pulchra 


ancestral 


181 
dorothea 
parowana 
fulgida 


! 

| : : 

! pimeriana 
| 


| 
miocene / pliocene 


pleistocene 


Figure 3. Hypothetical chronology of the four species in the C. pulchra group of the subgenus Cicindela. 


that the male genitalia, while remarkably similar, 
have constant specific differences. How and when 
C. pimeriana came to be in this area is unknown. 
It could have arrived from the north or from the 
east as an offshoot of C. fulgida. Its relationship 
with other species of the ‘pulchra’ group is of 
ancient origin, one that may be hypothesized as 
shown in Fig. 3. 

Some species of the Sonoran Cicindela fauna 
are so isolated and distinct as to suggest the pos- 
sibility that a region in southern Arizona and 
northern Sonora contained a specialized fauna 
since earliest Cenozoic times. These distinctive 
species are now reduced to relatively small areas, 
such as C. pimeriana in the San Bernardino and 
Sulphur Springs valleys, and C. beneshi and C. 
rockefelleri from the Gulf of California coast near 
Puerto Penasco. Though isolated and distinct, all 
three have related species within close range. 


LITERATURE CITED 


Casey, T. L. 1924. Memoirs on the Coleoptera, XI. Lancaster, 
PA (Privately published). 


Cazier, M. A. 1936. A review of the Willistoni, Fulgida, Paro- 
wana, and Senilis groups of the genus Cicindela. Bull. 
South. Calif. Acad. Sci. 35(Pt. 3): 156-163. 

- 1954. A review of the Mexican tiger beetles. Bull. 

Amer. Mus. Nat. Hist. 103(Art. 3): 231-309. 


Harris, E. D. 1913. Three new cicindelids. Jour. New York 
Entomol. Soc. 21(1): 67-69. 


Hubbs, C. L. and R. R. Miller. 1948. The zoological evi- 
dence/Correlation between fish distribution and hydro- 
graphic history in the desert basins of Western United 
States. In The Great Basin, with emphasis on Glacial and 
Postglacial times. Bull. Univ. Utah 38(20)(Biol. Ser. 10): 
17-166, figs. 10-29, 1 map. 


Knaus, W. 1922. Two new forms of Cicindela with remarks 
on other forms. Jour. New York Entomol. Soc. 30(4): 
194-197. 


Lawton, L. K. and H. L. Willis. 1974. Notes on the cicindelid 
collecting in the southwestern United States, summer 
1971. Cicindela 6(3): 44-65. 


Leconte, J. L. 1875. Notes on Cicindelidae of the United 
States. Trans. Amer. Entomol. Soc. 5: 157-162. 

McKee, E. D., R. F. Wilson, W. J. Breed and C. S. Breed, 
editors. 1967. Evolution of the Colorado River in Ari- 
zona. An hypothesis developed at the Symposium on 
Cenozoic Geology of the Colorado Plateau in Arizona, 
August 1964. Mus. North. Ariz., Bull. 44: i-ix, 1-67, figs. 
1-23, 1 map. 


Meinzer, O. E. and F. C. Kelton. 1913. Geology and water 
resources of the Sulphur Springs Valley, Arizona. United 
States Geological Survey, Water-Supply Paper 320: 1-231. 


Rivalier, E. 1954. Démembrement du genre Cicindela Linné, 
Il. Faune Ameéricaine. Rev. Fr. Entomol. 17(Fasc. 4): 249- 
268. 


Rumpp, N. L. 1957. Notes on the Cicindela praetextata- 
californica tiger beetle complex. Description of a new 
subspecies from Death Valley, California. Bull. South. 
Calif. Acad. Sci. 58(Pt. 3): 144-154. 


. 1961. Three new tiger beetles of the genus Cicindela 
from southwestern United States. Bull. South. Calif. 
Acad. Sci. 60(Pt. 3): 165-187. 


Willis, H. L. 1967. Bionomics and zoogeography of tiger 
beetles of saline habitats in the central United States. 
Univ. Kansas Sci. Bull. 47(5): 145-313. 


PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 5, pp. 183-213; 74 figs.; 1 table 


August 18, 1977 


A REVIEW OF THE NORTH AMERICAN GENERA OF LAPHYSTIINI, 
WITH A REVISION OF THE GENUS ZABROPS HULL 
(INSECTA: DIPTERA: ASILIDAE) 


By 


Eric Martin Fisher 


Department of Biology, California State University 
Long Beach, CA 90840 


ABSTRACT: The anatomical characteristics of the laphystiin genera Laphystia, Psilocurus, Perasis, and Zab- 
rops were studied, with emphasis placed on the male and female genitalia. Similarities in the genitalia of the 
Laphystiini and the Laphriinae provide evidence that these groups should be included together; structural 
differences in the genitalia were used to distinguish the genera studied. The following species and subspecies 
of Zabrops are recognized: flavipilis (Jones), tagax tagax (Williston), tagax argutus new subspecies, wilcoxi 
wilcoxi new species and subspecies, wilcoxi playalis new subspecies, wilcoxi arroyalis new subspecies, 
thologaster new species, and janiceae new species. Keys and illustrations are provided. 


INTRODUCTION 


The asilid tribe Laphystiini, nearly world- 
wide in distribution, is represented by four gen- 
era in North America: Laphystia Loew, 
Psilocurus Loew, Perasis Hermann, and Zab- 
rops Hull. Many genera in the tribe are poorly 
known taxonomically needing redefinition or re- 
vision. Concurrently new characters are needed 
to establish an improved classification of the 
tribe. 

The small genus Zabrops is especially in need 
of such revisionary work. These robber flies are 
commonly found during the spring and summer 
in the southwestern and central United States 
and northwestern Mexico. Although numerous 
specimens exist in collections, most are uniden- 
tified; only four specimens have been recorded 
in the literature, the last one in 1909. The iden- 
tity of the two species currently placed in the 
genus is obscure and several undescribed 
species are on hand. 


The purpose of this study is to provide de- 
scriptions of the taxa of Zabrops and to evaluate 
and illustrate new differentiating features of the 
North American genera of Laphystiini. 


MATERIALS 


This study is the result of the examination of 
1,025 adult specimens of Zabrops and approxi- 
mately 150 specimens of 60 species in 10 other 
genera of Laphystiini. 

The following abbreviations indicate the 
museums and private collections from which 
specimens were borrowed or examined: 
ASU—Arizona_ State University, Tempe, 
Arizona (M. Cazier); CAS—California Academy 
of Sciences, San Francisco, California (P. H. 
Arnaud, Jr.); CDA—Bureau of Entomology, 
California Department of Agriculture, Sac- 
ramento, California; CIS—California Insect 
Survey, University of California, Berkeley, 
California (E. I. Schlinger, J. Powell); CM—C. 


[183] 


184 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


H. Martin, Tucson, Arizona; CSLB—California 
State University, Long Beach (E. L. Sleeper); 
EF—E. M. Fisher, Long Beach, California; 
FSCA—Florida State Collection of Arthropods, 
Gainesville, Florida (H. V. Weems, Jr.); JW—J. 
Wilcox, Anaheim, California; LACM—Natural 
History Museum of Los Angeles County, Los 
Angeles, California (C. L. Hogue, R. R. Snel- 
ling); OSU—Ohio State University, Columbus, 
Ohio (C. A. Triplehorn); UCD—University of 
California, Davis, California (R. O. Schuster); 
UCR—University of California, Riverside, 
California (S. Frommer, J. D. Pinto); UK— 
Snow Entomological Museum, University of 
Kansas, Lawrence, Kansas (G. W. Byers); 
USNM—Smithsonian Institution, Washington, 
D.C. (L. V. Knutson). 


METHODS 


DissECTIONS.—Male and female genitalia 
were examined in from one to several represen- 
tatives of each population, depending on the 
number of specimens available. These tech- 
niques were used: (1) specimens to be dissected 
were placed in a relaxing chamber for periods of 
four to six hours in order to partially soften the 
tissues; (2) the apical half of the abdomen was 
cut off and placed in 10 percent KOH, heated to 
near boiling, for about 30 minutes; (3) the treated 
parts were washed in distilled water and excess 
tissue was removed; (4) the parts were then 
transferred to a watchglass with glycerine and 
dissected as necessary; (5) in males the 
hypopygium was removed from the abdomen, 
the epandrium separated from the gonopods, 
and the aedeagus removed; (6) for illustrative 
purposes, in some males the tergites and ster- 
nites 7 and 8, the dististyli, and the lateral pro- 
cesses were removed as well; (7) in females the 
tergite and sternite 8 were separated from the 
remnants of the abdomen; (8) after study, the 
parts were stored in glycerine in polyethylene 
microvials with silicone stoppers and the vials 
were placed on the pin with the specimen. 

ILLUSTRATIONS.—AIl drawings were made 
with the aid of a camera lucida attached to a 
stereomicroscope. 

Genitalia to be illustrated were placed in a de- 
pression slide on a minute spot of ‘‘Gelva”’ 
polyvinyl alcohol (pva), quickly positioned, then 
covered with glycerine and a cover glass. The 
pva prevented the structures from drifting while 


being drawn and had the added advantage of 
being easily dissolved with alcohol. 

The gonopods were slightly elevated an- 
teriorly when drawn in order to place the median 
process and the hypandrium in a _ horizontal 
plane. All bristles and hairs were omitted from 
the drawings. 

MEASUREMENTS.—AIl measurements were 
made with an ocular micrometer as follows: (1) 
width of face—measured immediately below the 
antennae; (2) width of eye—measured on same 
plane as face; both eyes were measured and an 
average taken; (3) length of body—measured 
from the anterior edge of the face to the tip of 
tergite 6; if the abdomen was curved (the usual 
case) the measurements were made on more 
than one plane for greater accuracy; (4) length of 
wing—taken from the base at the thorax to the 
tip; (5) width of tergite 3—measured from the 
dorsal view (can be interpreted as the width of 
the abdomen). Measurements | and 2 were made 
at 50x and measurements 3 to 5 at 12x. 

The means and standard errors of the above 
measurements, plus the ratios of eye/face widths 
and body/wing lengths, are listed in Table | for 
seven of the eight taxa of Zabrops. For Z. 


flavipilis (Jones), with only two specimens 


available, these measurements are given with 
the description. 

TERMINOLOGY .—In the descriptions of taxa, I 
have used terms as defined in Torre-Bueno 
(1962), with several exceptions. 

I have followed Karl (1959) for the terminol- 
ogy of nearly all the structures of the male 
genitalia and Martin (1968) for certain parts of 
the aedeagus. Crampton (1942) and Artigas 
(1971) were the sources of terms used for the 
female genitalia. 

The names of the lateral sclerites of the thorax 
are after Bonhag (1949), with a few differences. 
In the generic descriptions I used ANTERIOR 
BAND and POSTERIOR SPOT for areas on the 
pleural region that are usually bare of tomen- 
tum. The anterior band is a vertical area that 
extends from about the middle of the meso- 
anepisternum, down across the mesokatepister- 
num, to the base of the coxae. The posterior 
spot occupies most of the anterior three-fourths 
of the meron, just above and behind the middle 
coxae. 

PILE is used loosely and refers to any type of 
‘softer’ hair. TOMENTUM is the micro- 
pubescence of the integument and is used in- 


185 


FISHER: NORTH AMERICAN LAPHYSTIINI 


LOW Estat 
10 + IST 
COmsaSeul 
10° + SPI 
10° + 6F'1 
LOM arconl 
10° + Ly 
TOM Sal 
10° + 9F'I 
co” + LST 
10’ + Pr 
Oy) = (4S 1 
10 + Wil 
10° + PSI 
yysua] Sum 


/yigus] Apog 


10° = 97 
10’ = €€1 
ZO" + 6I'I 
10m tee 
Com aGal 
10° = v7 
10° = €7'l 
10° = 67 
Op neta 
10° = LUI 
ZO’ + 6l'l 
10° = 87 
10° + 97 
LO areal 
YIPIM 2905 
suipim akq 


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a) 
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Vale) 
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ND 
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aa 


Way (sg) 
os 
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NN 


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a 
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ma Om 
am 
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= 
2 
+ 


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06'1 


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= 
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LO’ + 66'S 
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stl eat 


D 
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10° 
10° 
10° 
10° 
C0 
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10° 
10° 
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10° 


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+| 


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qyipim 


690 10° + L8°0 (81 =N) 64 
L9°0 10° + 68°0 (Ol =N) Pe 
£9°0 10° + sZ°0 (Ol =N) 66 
19°0 10° + SZ°0 (Sc = N) PP 
9L'0 10° + 760 (€l =N) 66 
ILO 10° + 68°0 (LI = N) 22 
9L'0 10° + £60 (SC =N) 54 
0L'0 10° + 16°0 (Ss? = N) PP 
9L°0 10° + 98°0 (Ol =N) 36 
€L°0 10° + 98°0 (ZL) = IN) {242 
9L°0 10° + 16°0 (9 =N) 64 
cL'0 10° + 60 (tI = N) 22 
ILO 10° + 68°0 (SC =N) 36 
690 10’ + 06°0 (c¢ = N) 22 
20 yIpIm dAq 


(sojeduy soy eryeg) 
avaoiuol *7 


(unuInd ueg eBIyeg) 
Ia}Seso[OU} *Z 


(19[WIWIS) 
SIDAOMAD “MK *Z 


(yoeag OWSIq) 
syvanjd ‘Mm "7 


(o19RId “W) 
IXOJJIM “MK *Z 


(oursoy [A) 
SNINBAD XD8D] ‘7 


(OWA X ) 
xDdD] XDSD] ‘7 


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jo saisadsqns 2 saisedg 


“SdIDadSANS GNV SdIOdd§ sdoiqvZ dO SOILVY AGNV SLNAWAUNSVAPF GALOATAS AO (WW NI) SHOWUY GUVGNVLS GNV SNVAJY ‘[ ATAV 


186 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


stead of “‘pollen.’’ The latter term implies a 
dust-like or granular nature for this vestiture, 
which is not the case. Under very high magnifi- 
cation, tomentum is seen to consist of minute, 
recumbent, curved hairs or setae. In discussing 
the condition of pile on the mesonotum, I use 
‘‘anteriorly’’ to mean the entire anterior half of 
the disc, between the transverse suture and the 
anterior margin. 

CRITERIA USED FOR SPECIES AND SUBSPE- 
clEs.—AIl of the taxa that I have recognized in 
the genus Zabrops are allopatric—based on 
present knowledge of their distribution. This 
situation has led to some difficulty in defining 
them by the biological species concept, where 
reproductive isolation (most easily discerned in 
sympatric populations) is the main criterion 
(Mayr 1963). 

Therefore, it has been necessary to infer 
speciation (reproductive isolation) by differ- 
ences in the phenotype of populations or groups 
of populations. I have done this in the following 
manner: species have fundamental differences in 
the structure of the male genitalia, a usually 
greater overall differentiation of characters, and 
the absence of any intermediate character 
states; subspecies have no fundamental differ- 
ences in the male genitalia, usually less overall 
character differentiation, usually have intergra- 
dation of characters, and are geographically de- 
limited. I believe subspecies are of taxonomic 
value in Zabrops because they depict distinctive 
populations which are phenotypically and geo- 
graphically concordant. 

Variant populations not fitting the above 
criteria for a subspecies are discussed under 
their respective taxa. 


ACKNOWLEDGMENTS 


I am grateful for assistance from many people. 
Most especially, I thank Mr. Joseph Wilcox for 
the great amount of help, encouragement, and 
advice that he has given me on this project and 
on my study of robber flies in general. Indeed, 
most of the Zabrops specimens used in this 
study were loaned to me by Mr. Wilcox, who 
had either collected the material himself or had 
borrowed it from various museums. 

Mr. R. L. Westcott, Drs. R. C. Penrose, E. L. 
Sleeper, G. Mansfield-Jones, W. D. Stockton, 
and R. B. Loomis have all kindly reviewed this 
paper and offered many helpful suggestions. 

Thanks also to the late Dr. Charles H. Martin 


for the numerous discussions we have had on 
the taxonomy of Diptera and for a translation of 
the valuable paper by Karl, and Dr. Paul H. Ar- 
naud, Jr. for the many kindnesses he has ex- 
tended me on visits to the California Academy of 
Sciences. 

I owe much gratitude to the people listed in 
the materials section for the loan of specimens 
used in this study, but I especially thank Dr. 
George W. Byers for sending me the holotype of 
Zabrops tagax to examine. 

Finally, I give very special thanks to my wife 
Janice for her help in many aspects of this study. 


TAXONOMIC SECTION 
Tribe LAPHYSTIINI 


CHARACTERISTICS AND RELATIONSHIPS.— 
The tribe Laphystiini consists of about 25 de- 
scribed genera and is found in all the faunal re- 
gions, but the Ethiopian, southern Palearctic, 
and Neotropical are best represented. One 
genus each occurs in the Oriental and Australian 
regions, while four genera are found in the 
Nearctic (Hull 1962). 

The taxonomic position of the Laphystiini has 
been somewhat controversial, various authors 
placing them in the subfamily Dasypogoninae, 
the subfamily Laphriinae, or in the Dasy- 
pogoninae with the Laphriinae also included. 
Wilcox (1960) and Oldroyd (1963) reviewed the 
taxonomic history of the Laphystiini, Oldroyd 
concluding that the tribe belonged in the 
Laphriinae. Oldroyd (1970) subsequently in- 
cluded the genera of Laphystiini in the tribe 
Laphriini. 

I believe that the Laphystiini is indeed closely 
related to the Laphriini and should be placed in 
the same subfamily with that group but with 
separate tribal status. The primary evidence for 
this close relationship is the great similarity in 
structure between male and female genitalia of 
both these groups. Karl (1959), in a comprehen- 
sive study of the male genitalia of Asilidae, was 
the first to mention this similarity. He compared 
the male genitalia of Hoplistomerus nobilis 
Loew, a laphystiin from Africa, with those of 
three genera and nine species of Laphriini and 
concluded: *‘The single principal difference con- 
sists only in that Hoplistomerus has still no 
hypopygial inversion’’ (translated). Even this 
difference can be eliminated because Karl was in 
error in believing that the hypopygium of Hop- 


FISHER: NORTH AMERICAN LAPHYSTIINI 


listomerus is not inverted. I have examined six 
male specimens of H. nobilis and one male each 
of H. engeli Oldroyd and H. zelimina Speiser; in 
all specimens the genitalia was rotated 180°. Ap- 
parently the single specimen of H. nobilis that 
Karl studied was collected before rotation (in- 
version) had occurred. 

The general structure of the laphystiin 
hypopygium is as follows (partly from Karl 
1959): (1) the hypandrium is reduced or absent 
(in which case it 1s possibly “‘absorbed”’ in the 
coalescing of the basistyli); (2) the gonopods 
have large basistyli, fused medially; (3) the 
basistyli uniformly have two pairs of append- 
ages, the dististyli and the lateral processes— 
these structures assume a wide variety of shapes 
within the tribe and are important as taxonomic 
characters; (4) the epandrium is coalesced into a 
single, cupped, shield-like structure; (5) the ven- 
tral lamellae are well developed and are longer 
than the cerci; (6) the aedeagus has three tubes, 
one dorsal and two ventral, which are partially 
to entirely coalesced; (7) the hypopygium is ro- 
tated from 90° to 180° around the longitudinal 
axis of the abdomen. 

The Laphystiini are further characterized by a 
great size reduction of abdominal segments 7 
and 8. In males of this tribe only tergites | to 6 
are visible from above; the remaining tergites 7 
and 8 are hidden beneath the much larger tergite 
6. Tergite 8 is reduced to a fraction of the length 
of tergite 6, while tergite 7 is not quite so re- 
duced in length; sternite 8 is reduced in width, 
assuming a rounded or rhomboid shape; sternite 
7 is either entirely absent or is represented as a 
minute vestigial sclerite. Karl (1959) attributes 
this size reduction to the rotation of the 
hypopygium, segments 7 and 8 being partly ro- 
tated also, with segment 8 affected the most. 
This reduction is evident in other asilids with 
inverted genitalia (e.g., other Laphriinae and 
certain genera of Dasypogoninae) but is greatest 
in this tribe. 

In the female genitalia, sternite 8 generally has 
a subapical depression and from one to three 
apical notches. Acanthophorites are absent, but 
frequently there are strong bristles apically on 
tergites 8 or 9, or on sternite 8. The spermatheca 
consists of three long filamentous tubes, the dis- 
tal half of each loosely coiled. The tubes empty 
into the bursa which is supported by a lightly 
sclerotized, U-shaped furca. The spermathecae 
are remarkably uniform in appearance in the 


187 


Laphystiini, at least among the North American 
genera. 

In the wings of the Laphystiini, the marginal 
cell is open to narrowly closed; when closed, the 
second vein (R,,3) is usually recurrent. The first 
posterior cell is open to closed and petiolate; the 
fourth posterior and anal cells are closed and 
usually petiolate. In many genera the costa is 
reduced (in width and/or length) or absent be- 
yond the apex of the wing. The other groups of 
Laphriinae have a similar venation but the mar- 
ginal cell is always closed and long-petiolate and 
the second vein is never recurrent. 

The head is relatively broad, averaging about 
one and one-half to one and three-fourths times 
wide as high. The antennae are always provided 
with an apical pit enclosing a small spine. Usu- 
ally this pit is preceded by a one- or two- 
segmented microsegment (the style); at least one 
genus, Psilocurus, lacks the style. The palpi are 
two-segmented. 

In the Laphystiini, the prosternum is com- 
plete, fused with the pronotum laterally, as in 
the majority of asilids. A mesopleural bristle is 
sometimes present (e.g., in some species of 
Hoplistomerus, Trichardis and Martinia, at 
least) but usually is absent; the presence of this 
bristle has been used to characterize the Laph- 
riinae in the past. The postscutellar slopes 
(mediotergite) are micropubescent—never with 
hair or bristles as in some other Laphriinae. The 
postmetacoxal area is membranous. 

In some genera of laphystiins, the hind femora 
is strongly swollen and with ventral, setiferous 
tubercules. The anterior and middle tibiae are 
without any modified bristles or spines. The 
pulvilli are occasionally abbreviated, or even 
absent, in certain genera. 

The adults are characteristic inhabitants of 
open areas where they are found resting either 
directly on the ground or on pebbles or decum- 
bent herbaceous vegetation very close to the 
ground. Laphystiini are commonly encountered 
in such habitats as alkaline flats, sand dunes, sea 
beaches, muddy or dry margins of streams, ar- 
royos, fields, roadsides, and paths in more 
forested areas. Little is known about the biol- 
ogy of the immature stages of laphystiins. 
Krivosheina (1973:459) found that larvae and 
pupae of the Palearctic species Laphystia 
carnea Hermann ‘‘develop in sandy loam soils, 
usually by edges of depressions, in regions with 
tamarisk and various annual saltworts.”’ It 


188 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


seems likely that the other genera of this tribe 
are also soil dwelling because the adults are 
often found in locales where suitable wood is 
scarce. The other groups of Laphriinae are pre- 
dominately forest inhabiting, with adults which 
mainly sit on leaves, branches, or logs; as far as 
is known, their larvae live in decaying wood 
where they are predaceous on the immature 
forms of wood-burrowing Coleoptera and Hy- 
menoptera (Hull 1962; Knutson 1972). 

Many of the aforementioned attributes are not 
individually exclusive to the Laphystiini. How- 
ever, when taken as a whole, they serve to 
characterize the group well. Also, the ecological 
differentiation between this tribe and the re- 
mainder of the Laphriinae, together with the 
anatomical differences in the male abdomen and 
the wing venation, justify keeping the Laphys- 
tiini as a separate tribe. 

THE NorRTH AMERICAN GENERA OF 
LAPHYSTIINI.—Iwo keys, published recently, 
serve to separate the four genera of North Amer- 
ican Laphystiini: Wilcox (1960), for the United 
States, and Hull (1962), for the world. However, 
with the discovery of additional species and 
variation in Zabrops, along with a wider varia- 
tion of characters in other genera, a revised key 
is desirable. Such a key is presented below, fol- 
lowed by diagnostic descriptions of the genera. 

A fifth North American genus, Bohartia Hull, 
was included by Hull (1962) in the Laphystiini 
but was subsequently transferred to the Dioct- 
riini by Martin and Wilcox (1965). These charac- 
teristics indicate that Bohartia is correctly 
placed in the Dioctriini: wing venation with all 
cells open; prosternum reduced—isolated later- 
ally from the pronotum; males with gonopods 
divided and with abdominal segments 7 and 8 
not reduced, visible from above. 


KEY TO GENERA OF NORTH AMERICAN LAPHYSTIINI 


la. Face gibbose on lower 70 percent or less, narrower 
thaney.cmees sees = 5 Boo See ee oe eee 2 

lb. Face entirely gibbose, equal to 1.2 times width of eye; 

(first posterior cell usually open; hypandrium absent; 

abdominal tergites 1 to 6 with lateral bristles) ______ 
Se ee Sie be Laphystia Loew 


2a. First posterior cell closed; face wide, 60 to 90 percent 
width of eye; antennal style present; hypandrium 
present 

2b. First posterior cell widely open; face narrower, 50 to 
60 percent width of eye; antennal style absent; hy- 
pandrium absent; (abdominal tergites 1 to 6 with lat- 
eralibristless)haee ean ee eee Psilocurus Loew 


3a. Face most prominent at oral margin, gradually receding 
above; mystax with a row of strong bristles on oral 
margin and dense, recumbent, squamose hairs above; 
lateral bristles present on abdominal tergite 1 only; 
pronotum with strong dorsal and posterolateral bris- 
HES ean 2 ole oe ee oe Perasis Hermann 

3b. Face evenly gibbose on more than lower half; mystax 
consisting of long, slender bristles over entire gib- 
bosity; abdominal tergites 1 and 2 up to | through 5 
with lateral bristles; pronotum with pile only________ 
ee ee en eae) ee ES Zabrops Hull 


Genus Laphystia Loew 


Laphystia Loew, 1847: 538. [Type-species: sabulicola Loew 
(monotypy).] 


DESCRIPTION.—Small to medium sized flies 
with short pile and robust form; length 7 to 13 
mm. 

Head (Fig. 1): face wide, at antennae subequal 
to one and one-fourth times width of eye, 
slightly wider at front, evenly produced from 
oral margin to antennae; gibbosity covered with 
pile, longest and densest at oral margin where 
there usually are some slender bristles. Front, 
vertex, and ocellar tubercule with abundant pile; 
occiput with both slender bristles and pile; beard 
thick and rather long. Proboscis short, some- 
what robust; palpi small, second segment with a 
few fine hairs. 

Antennae with segment | subequal to one and 
one-half times as long as segment 2, segment 3 
subequal to one and one-third length of seg- 
ments | and 2 together; segment 3 slightly swol- 
len toward middle; sensory area variable, oc- 
cupying apical one third to two thirds of inner 
side; style two segmented, the apical segment 
much longer than the basal and with an oblique 
excavation bearing a small spine; segment | with 
two strong ventral bristles and numerous hairs 
only. 

Thorax: pronotum with hairs only; meso- 
notum evenly covered with pile, which var- 
ies from short and recumbent to longer and 
erect; dorsocentral bristles absent; two to three 
notopleural, supraalar, and postalar bristles 
present. Scutellum with marginal hairs or bris- 
tles, length and number variable. Pleura evenly 
tomentose, without polished areas; lateroter- 
gites with numerous long, slender bristles. 

Legs: femora with diameter slightly greater 
than that of tibiae, and with numerous hairs and 
several dorsoapical bristles; tibiae and tarsi with 
bristles longer and more abundant; pulvilli vary- 


FISHER: NORTH AMERICAN LAPHYSTIINI 189 


—~ 
SSS 
, SS 


= 


ASR 

ZR 
ans NAS 
V fi ) AN ARS EO \ 


SSS 
= 
SS = 
SS 
SSS 


SS 


—— 
— 


— 


2.0 iit ————————| 


oY 


We x 
ZIAPNSN 


LI 


ae 


ae 


Ficures | to 4. Heads of Laphystiini, lateral view. Fig. 1. Laphystia jamesi. Fig. 2. Psilocurus sp. Fig. 3. Perasis argentifacies. 


Fig. 4. Zabrops tagax tagax. 


190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


ing from well developed to abbreviated to nearly 
absent. 

Wings: marginal cell narrowly closed to nar- 
rowly open; first posterior cell generally open, 
but closed and petiolate in several species; 
fourth posterior and anal cells closed petiolate; 
costa extending to apex of anal cell, absent be- 
yond or, rarely, ending at apex of wing. 

Abdomen (Figs. 10 to 12): pile very short and 
recumbent dorsally, longer and erect laterally 
and ventrally in some species; tergites | to 6 
with two or more strong lateral bristles. Males 
with visible portion of tergite 6 about as long as 
tergite 5; tergite 7 with middle three-fourths re- 
duced in length; tergite 8 greatly reduced in 
length save for a posteromedian lobe; sternite 7 
absent. 

Genitalia, male (Figs. 5 to 9): hypandrium ab- 
sent. Gonopods with basistyli semi-membranous 
along midline (incompletely fused?); dististyli 
simple, like an elongate lobe; lateral processes 
narrow, with apex semiacute and slightly re- 
curved. Aedeagus with basal part and ejacula- 
tory apodeme moderate sized; tubes coalesced 
and quite thin apically, curving dorsad. Epan- 
drium wider than long, ventral lamellae and 
cerci slightly emarginate apically. Hypopygium 
without prominent bristles. 

Genitalia, female (Fig. 13): sternite 8 with 
apex with three notches, the median most pro- 
nounced, forming four lobes; tergite 8 with sev- 
eral to numerous slender, apical bristles. Furca 
of spermatheca either split or entire basally. 

REMARKS.—The genus Laphystia contains 
about 55 described species, the majority from 
the Northern Hemisphere; these are about 
equally divided between the Old World and the 
New World. Probably most, if not all, of the 
species described from south of the equator as 
Laphystia belong in other genera. 

Wilcox (1960) has revised the North American 
species of Laphystia. 


Genus Psilocurus Loew 


Psilocurus Loew, 1874: 373. [Type-species: nudiusculus Loew 
(monotypy).] 


Orthoneuromyia Williston, 1893: 67. [Type-species: modesta 
Williston (monotypy).] 


DESCRIPTION.—Small sized, slender flies with 
reduced pile; 6 to 13 mm long. 

Head (Fig. 2): face narrow, at antennae about 
50 to 60 percent as wide as eye, noticeably 
wider at frons; lower 45 to 65 percent gibbose 
and covered with strong bristles, remainder 
plane and with short hairs; sometimes a small 
swelling is present immediately below antennae. 
Front and vertex with a few scattered short 
hairs, ocellar tubercule bare. Occiput with a row 
of strong bristles; beard short pilose. Proboscis a 
little longer than face; palpi moderate sized, 
segment 2 spindle-shaped and with several long 
slender bristles. 

Antennae: segment | slightly longer than seg- 
ment 2, together subequal to segment 3; segment 
3 generally strongly swollen at middle, tapering 
towards ends; sensory area occupying apical 
one half or less; style absent, apex of segment 3 
with a broad oblique concavity bearing a small 
spine; segment | ventrally with a strong bristle 
and scattered bristly hairs, segment 2 with only 
bristly hairs. 

Thorax: pronotum with a double row of fine 
bristles, laterally with pile. Mesonotum with 
dense, very short, recumbent setate pile; dorso- 
central bristles greatly reduced or absent; one 
notopleural, one supraalar and one _ postalar 
bristle—all strong. Scutellum with two to four 
strong marginal bristles. Pleura with bands and 
spots subshining to polished; laterotergite with 
five to seven slender bristles and some shorter 
hairs. 

Legs: femora somewhat swollen, about half 
again as wide as tibiae, with one to three subapi- 
cal dorsal bristles; tibiae and tarsi with bristles 
more numerous apically; pulvilli well developed. 

Wings: marginal and first posterior cells wide 
open; fourth posterior and anal cells closed and 
short petiolate; costa well developed, complete. 

Abdomen (Figs. 19 to 21): pile reduced as on 
mesonotum. Tergite 1 with two to five strong 
bristles laterally; tergites 2 to 6 each with one to 


=> 


Figures 5 to 13. Terminal abdominal segments and genitalia of Laphystia spp. (Figs. 5 to 12 Laphystia jamesi males, Fig. 13 
Laphystia litoralis Curran female.) Fig. 5. Gonopods and aedeagus, ventral view, dististyli horizontally lined: ae.-aedeagus; 
dist.-dististylus; |.p.-lateral process; bst.-basistylus. Fig. 6. Dististylus, lateral view, dorsal surface on left. Fig. 7. Lateral process, 
lateral view, dorsal surface on left. Fig. 8. Epandrium, dorsal view: v.l.-ventral lamellae; ce.-cerci; ep.-epandrium. Fig. 9. 
Aedeagus, lateral view, ventral surface above: e.a.-ejaculatory apodeme; b.p.-basal part; tu.-tubes. Fig. 10. Sternite 8, ventral 
view. Fig. 11. Tergite 8, dorsal view. Fig. 12. Tergite 7, dorsal view. Fig. 13. Ventral view, sternite 8 vertically lined: t.9-tergite 9; 


ce.-cerci; t.8-tergite 8; s.8-sternite 8; fu.-furca of spermatheca. 


19] 


FISHER: NORTH AMERICAN LAPHYSTIINI 


192 


two strong bristles laterally. Males with visible 
portion of tergite 6 longer than tergite 5; tergite 7 
with a posteromedian T-shaped projection; ter- 
gite 8 quite short, trilobed posteriorly; sternite 7 
absent. 

Genitalia, male (Figs. 14 to 18): hypandrium 
absent. Gonopods with dististyli bearing a wide, 
apically rounded dorsal lobe and a posteriorly 
directed, ventral, finger-like projection; lateral 
processes long and tapered, projecting beyond 
apex of genitalia in most species. Aedeagus with 
basal part well developed and “‘box-like’’; tubes 
trifurcate distally, with one dorsal and two ven- 
tral parts; ejaculatory apodeme small, produced 
ventrad. Epandrium about as wide as long; ven- 
tral lamellae well developed, produced into two 
apically tapering, strongly sclerotized lobes; 
cerci short, with median notch. Epandrium with 
a few posterolateral bristles. 

Genitalia, female (Fig. 22): sternite 8 rounded 
apically, with shallow median notch; central part 
excavated. Tergite 8 with several apical bristles. 
Spermatheca with furca divided basally. 

REMARKS.—Nine species of Psilocurus have 
been described from the United States, Mexico, 
and Ecuador (Hull 1962), and two species are 
known from western Asia (Lehr 1969). Numer- 
ous species remain undescribed, especially from 
Mexico and Central America. 


Genus Perasis Hermann 


Perasis Hermann, 1905: 37. [Type-species: sareptana Her- 
mann (monotypy).] 

Triclis, of authors, not of Loew. 

Saucropogon Hull, 1962: 103. [Type-species: Perasis trans- 
vaalensis Ricardo (original designation). ] 


DESCRIPTION.—Small to medium _ sized, 
stout-bodied, black flies with greatly reduced 
pile; length 8 to 13 mm. 

Head (Fig. 3): face about two-thirds to sub- 
equal width of eye, slightly wider at frons, pro- 
duced at oral margin and gradually receding to 
about middle, upper one-half plane; oral margin 
with row of about 10 to 15 strong bristles; lower 
two-thirds of face densely covered with ap- 
pressed, squamose bristles, upper one-third with 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


short, erect pile. Front with many slender bris- 
tles laterally. Vertex with scattered hairs, ocel- 
lar tubercule with about 12 slender bristles. Oc- 
ciput with 25 to 30 strong bristles; beard sparse 
and short. Proboscis well developed, twice as 
long as face; palpi relatively large, segment 2 
with many bristles. 

Antennae: segment | one and one-half times 
as long as segment 2; segment 3 nearly one and 
one-half times as long as segments | and 2 to- 
gether, not or slightly swollen; sensory area 
large, occupying apical two-thirds of inner side 
of segment 3; style with one segment, about 
one-fourth as long and one-half as wide as seg- 
ment 2, broadly excavated apically and bearing a 
central spine; segments | and 2 each with sev- 
eral slender bristles. 

Thorax: pronotum with about 20 bristles dor- 
sally and 2 to 4 bristles laterally, all strong and 
spiniform. Mesonotum evenly covered with ex- 
tremely short, recumbent pile, mostly spinelike; 
dorsocentral bristles absent; one notopleural, 
two to three supraalar, and two to three postalar 
bristles, all very strong. Scutellum with recum- 
bent marginal hairs only. Pleura with anterior 
band and posterior spot bare of tomentum; 
laterotergite with six to eight strong bristles. 

Legs: femora not swollen, slightly wider than 
tibiae; bristles of legs stout but reduced in length 
and number. Pulvilli well developed. 

Wings: marginal cell open; first posterior, 
fourth posterior, and anal cells all closed and 
short petiolate; costa complete but greatly re- 
duced in thickness beyond apex of wing. 

Abdomen (Figs. 28 to 31): pile reduced as on 
mesonotum. Tergite | with four to six strong 
bristles laterally; remaining tergites without bris- 
tles. Males with sternite 7 present; tergites 7 and 
8 with middle one-third reduced in length. 

Genitalia, male (Figs. 23 to 27): hypandrium 
present, triangular, partially fused to basistyli. 
Gonopods with dististyli angulate in middle, the 
apex recurved; lateral processes slightly curved, 
tapering to apex. Aedeagus with basal part well 
developed and with a pair of secondary pro- 
cesses which extend posteriorly, paralleling 


— 


FiGures 14 to 22. Terminal abdominal segments and genitalia of Psilocurus sp. (Figs. 14 to 21 males, Fig. 22 female). Fig. 14. 
Gonopods and aedeagus, ventral view, dististyli horizontally lined. Fig. 15. Dististylus, lateral view, dorsal surface on left. Fig. 16. 
Lateral process, lateral view, dorsal surface on left. Fig. 17. Aedeagus, lateral view, ventral surface above. Fig. 18. Epandrium, 
dorsal view. Fig. 19. Sternite 8, ventral view. Fig. 20. Tergite 8, dorsal view. Fig. 21. Tergite 7, dorsal view. Fig. 22. Genitalia, 


ventral view, sternite 8 horizontally lined. 


194 PROCEEDINGS OF THE CALJFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


tubes; tubes relatively wide, coalesced distally; 
ejaculatory apodeme large. Epandrium about as 
wide as long; ventral lamellae with apex entire; 
cerci deeply emarginate apically. Gonopods 
with numerous posterolateral bristles. 

Genitalia, female (Fig. 32): apex of sternite 8 
with lateral notches large; median notch smaller, 
lobes approximate; area anterior to median lobes 
with pronounced concavity. Tergite 8 without 
bristles. Furca of spermatheca with base large. 

REMARKS.—Perasis Hermann is a primarily 
Old World genus with currently one New World 
species, argentifacies (Williston), from south- 
central Mexico. Originally described by Williston 
as a Triclis, argentifacies was provisionally 
placed in Perasis by Wilcox in 1960; Martin and 
Wilcox (1965) and Martin and Papavero (1970) 
have maintained this treatment. As pointed out 
by Oldroyd (1970), Hull (1962) confused the iden- 
tity of Perasis by presenting a description of that 
genus based on material misidentified as the 
type-species (sareptana Hermann) and possibly 
belonging to another genus. Furthermore, he 
erected a new genus Saucropogon (Hull 1962) as 
a result of his misinterpretation. Saucropogon is 
the same as Perasis (sensu Hermann) and was 
synonymized with that genus by Oldroyd (1970). 

Based on Hull’s thorough description of Sauc- 
ropogon (1.e., Perasis), | believe that Wilcox was 
correct in his placement of argentifacies. This 
species agrees in every detail with Hull’s descrip- 
tion and some of the characters, namely the 
structure of the mystax, face, and lateral pronotal 
bristles, seem to be diagnostic for the genus 
among the Laphystiini. Comparison of the male 
genitalia of both Old World and New World 
species would offer conclusive evidence; unfor- 
tunately I have not had the opportunity to 
examine Old World material. 

In addition to argentifacies, several unde- 
scribed species of Perasis exist in Mexico and the 
United States. 

Among the North American Laphystiini, 
Perasis is most similar to Zabrops. These genera 


share several characteristics: same wing vena- 
tion; possession of hypandrium and sternite 7 in 
males; similar type dististyli and female sternite 
8. Despite these similarities, many differences in 
other character states (cited in descriptions) indi- 
cate that these two genera are not especially 
closely related. 


Genus Zabrops Hull 


Zabrops Hull, 1957: 90. [Type-species: Triclis tagax Williston 
(original designation). ] 
Triclis, of authors, not of Loew. 


DESCRIPTION.—Small to medium sized flies 
with short pile, 6 to 11 mm long. 

Head (Fig. 4): face at antennae 60 to 90 percent 
width of eye, slightly wider below and at front; 
lower 55 to 70 percent gibbose and covered with 
slender bristles subequal in length to antennae; 
plane part with short hairs. Front with hairs and 
slender bristles laterally. Vertex with numerous 
hairs, ocellar tubercule with two or four bristles. 
Occiput with about 20 bristles, usually strong; 
beard moderately dense and long. Proboscis sub- 
equal to length of face; palpi small, segment 2 
with several long hairs. 

Antennae: segment | subequal to one and 
one-half times as long as segment 2; segment 3 
subequal to three-fourths as long as segments | 
and 2 together, slightly to strongly swollen just 
beyond middle; sensory area small, restricted to 
apical one-third of inner side; style one or two 
segmented, about one-third as long and wide as 
antennal segment 2, apex obliquely concave and 
with minute spine; segments | and 2 with numer- 
ous hairs, segment | with one to several strong 
bristles ventrally. 

Thorax: pronotum with hairs only. Meso- 
notum with pile sparse and recumbent to 
dense and erect; dorsocentral bristles hairlike, 
not or weakly differentiated from other dorsal 
pile; two to four notopleural, supraalar, and post- 
alar bristles. Scutellum with 10 to 20 long margi- 
nal bristles. Pleura with anterior band and pos- 


— 


FIGURES 23 to 32. Terminal abdominal segments and genitalia of Perasis argentifacies (Figs. 23 to 31 males, Fig. 32 female). Fig. 
23. Gonopods, hypandrium, and aedeagus, ventral view, dististyli horizontally lined, secondary process stippled: hy.-hypandrium 
(vertically lined). Fig. 24. Dististylus, lateral view, dorsal surface on left. Fig. 25. Lateral process, lateral view, dorsal surface on 
left. Fig. 26. Epandrium, dorsal view. Fig. 27. Aedeagus, lateral view, ventral surface above: s.p.-secondary process (stippled). 
Fig. 28. Sternite 8, ventral view. Fig. 29. Sternite 7, ventral view. Fig. 30. Tergite 8, dorsal view. Fig. 31. Tergite 7, dorsal view. Fig. 


32. Genitalia, ventral view, sternite 8 vertically lined. 


FISHER: NORTH AMERICAN LAPHYSTIINI 


i a ———— 


196 


terior spot bare of tomentum; laterotergite with 
about 20 long, slender bristles. 

Legs: femora swollen, nearly twice as wide as 
tibiae. Bristles of femora, tibiae and tarsi long, 
rather strong. Pulvilli well developed. 

Wings: marginal cell open; first posterior cell 
closed, usually petiolate; fourth posterior and 
anal cells closed and always petiolate. Costa 
complete, or greatly reduced in width (or absent) 
beyond apex of wing. 

Abdomen (Figs. 38 to 41): pile short and ap- 
pressed. Bristles numerous laterally on tergites 1 
and 2; tergite 3, and sometimes tergites 4 and 5, 
with one or two lateral bristles; tergite 6 with 
visible portion one-half to two-thirds as long as 
tergite 5; tergite 7 broad, tergite 8 much reduced 
in middle third; sternite 7 present. 

Genitalia, male (Figs. 33 to 37): hypandrium 
present, fused to basistyli of gonopods, bilobed, 
the lobes divergent to contiguous. Gonopods 
with a posteriorly directed median process which 
is bifurcate or entire; dististylus angulate in mid- 
dle, the apex acute and recurved; lateral pro- 
cesses strongly clavate. Aedeagus with basal part 
reduced; tubes somewhat wide, coalesced dis- 
tally; ejaculatory apodeme large. Epandrium 
longer than wide; ventral lamellae and cerci 
notched apically, not strongly sclerotized. Both 
gonopods and epandrium with numerous, strong, 
posterolateral bristles. 

Genitalia, female (Fig. 42): apex of sternite 8 
with lateral lobes large; median lobes feebly indi- 
cated by shallow emargination. Tergite 8 with 10 
or more strong black bristles apically. 

All species of Zabrops are sexually dimorphic. 
Besides the external differences in the genitalia of 
the two sexes, females have a wider face, longer 
wings, and wider abdomen—which generally is 
more extensively tomentose dorsally—than in 
the males. In addition, the hind tibiae are arcuate 
in males, straight in females. 

TAXONOMIC History .—Hull (1957) described 
the genus Zabrops, with Triclis tagax Williston 
(1883) as the type-species. He stated that the 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


other American Triclis (flavipilis Jones [as 
‘‘flavipes’’| and argentifacies Williston) proba- 
bly belonged to Zabrops also. Triclis tagax was 
originally described from a female specimen col- 
lected in Kern County, California. Williston 
(1886) later presented a brief note on a male 
specimen of tagax from an unknown locality. A 
second species of Triclis, argentifacies, was de- 
scribed by Williston (1901) from material col- 
lected in Guerrero, Mexico. 

Jones (1907) supplied the third North Ameri- 
can species, Triclis flavipilis, based on a single 
female from Meadow, Nebraska. He indicated 
that his new species was close to tagax and that 
Triclis should possibly be synonymized with the 
genus Laphystia. Back (1909) studied an addi- 
tional female specimen of tagax from California, 
commented on the possible synonymy of Triclis 
and Laphystia, and discussed several new 
characters for separating the two genera. 

In a revision of the North American species of 
Laphystia, Wilcox (1960) presented a key to the 
regional genera of Laphystiini, used Hull’s re- 
cently described genus Zabrops for the species 
of Triclis from the United States, and transferred 
the Mexican species argentifacies to the previ- 
ously strictly Old World genus Perasis Her- 
mann. Hull (1962) treated the world Asilidae, 
with descriptions and keys to the genera, includ- 
ing Laphystiini. Hull had not yet seen Wilcox’s 
work and repeated his statement that both 
flavipilis (as ‘‘flavipes’’) and argentifacies prob- 
ably belonged in Zabrops, although he formally 
listed them under Triclis. Martin and Wilcox 
(1965) listed tagax and flavipilis in Zabrops and 
argentifacies in Perasis. 

SPECIES AND SUBSPECIES OF ZABROPS.—In 
the following key to the taxa of Zabrops it is 
necessary to determine the shape of the median 
process of the gonopods to properly identify 
males of the tagax species group. This can be 
done without dissection—see Figs. 46 and 47. 
The median process is more heavily sclerotized 
than the hypandrial lobes and appears black, in 


= 


FIGURES 33 to 42. Terminal abdominal segments and genitalia of Zabrops tagax tagax (Figs. 33 to 41 males, Fig. 42 female). Fig. 
33. Gonopods and hypandrium, ventral view, dististyli horizontally lined, median process stippled, hypandrium vertically lined: 
m.p.-median process. Fig. 34. Dististylus, lateral view, dorsal surface on left. Fig. 35. Lateral process, lateral view, dorsal surface 
on left. Fig. 36. Epandrium, dorsal view. Fig. 37. Aedeagus, lateral view, ventral surface above. Fig. 38. Sternite 8, ventral view. 
Fig. 39. Sternite 7, ventral view. Fig. 40. Tergite 8, dorsal view. Fig. 41. Tergite 7, dorsal view. Fig. 42. Genitalia, ventral view, 


sternite 8 lined. 


198 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


contrast to the reddish-brown color of the latter 
structures. 


4b. 


Sa. 


Sb. 


6a. 


6b. 


as 


7b. 


8a. 


8b. 


9a. 
9b. 


KEY TO SPECIES AND SUBSPECIES OF ZABROPS 


. Front tomentose, with at most a small, shiny black 


spot above antennae; occipital bristles white to yel- 
lowish brown; males with hypandrial lobes well 
separated and rounded apically; southwestern 
United States and northwestern Mexico species ___ 2 


. Lower half of front shiny black; occipital bristles black; 


males with hypandrial lobes subcontinguous and 
apically emarginate; central United States species 
es ee a ae ee ee ee flavipilis (Jones) 


. Costa complete; males with median process of gono- 


pods entire or with apex notched; females with ex- 
tensive shiny areas on abdominal tergites; (tagax 


TOU P) eee a eee ee ee en eee ee 3 
. Costa evanescent beyond apex of wing; males with 
median process of gonopods completely bifid; fe- 
males with abdominal tergites all tomentose; (tholo- 
PUSTCTARLOUP) yee aie ee ee a ee ns VN 12 
5 WAGs ee ee ee ee ee ee 4 
Renal 6S a ee eee ee 8 


. Gonopods with median process short, wider than long, 


the apex notched; southern California, southern 
Arizona, northwestern Mexico species; (tagax (Wil- 
LStOn)) eee ee re ee 5 
Gonopods with median process longer, twice as long as 
wide, the apex rounded or truncate; central and 
northern California species; (wilcoxi new species) 6 


Anterior margin of abdominal tergite 1 tomentose; 
erect hairs of mesonotum densest in dorsocentral 
rows, sparse elsewhere; marginal bristles of scutel- 
lum partly yellow; Mojave and Sonoran desert sub- 
species tagax tagax (Williston) 

Anterior margin of abdominal tergite 1 bare of to- 
mentum; erect hairs of mesonotum dense, equally 
distributed over disc; marginal bristles of scutellum 
all black; coastal southern California and Baja Cali- 
fornia subspecies ____ tagax argutus new subspecies 


Humeral area densely tomentose; mystax entirely 
white or yellow 
Humeral area bare of tomentum; upper third of mystax 
black, remainder white 
wilcoxi playalis new subspecies 


Apical abdominal tergites red; scutellum with marginal 
bristles all black __ wilcoxi arroyalis new subspecies 

Apical abdominal tergites black; scutellum with mar- 
ginal bristles partly to entirely yellow ______________ 

wilcoxi wilcoxi new subspecies 


Abdominal tergites 2 to 4 with posterior tomentose 
fasciae interrupted by one-third their width or more; 
tergite 1 with broad posteromedian bare area 

Abdominal tergites 2 to 4 with posterior tomentose 
fasciae complete, or interrupted at middle by less 
than one-fourth their width; tergite 1 tomentose, with 
a small posteromedian bare spot ________-_________ 

doackas Sala a Phe ne Ne tagax tagax (Williston) 


Apical abdominal tergites mostly or entirely black __ 10 
Apical abdominal tergites mostly red _______________ 11 


10a. Mystax, erect mesonotal hairs and scutellar bristles 
alliblack: = _- == tagax argutus new subspecies 

Mystax yellow; erect mesonotal hairs and scutellar 
bristles partly or entirely yellow 


wilcoxi wilcoxi new subspecies 


10b. 


lla. Humeral area bare of tomentum; abdominal tergites 
2 to 4 with posterior fasciae interrupted by about 
two-thirds the width of abdomen ___---- 

wilcoxi playalis new subspecies 
11b. Humeral area densely tomentose; abdominal tergites 
2 to 4 with posterior fasciae interrupted by about 


one-third the width of abdomen __________-__________ 


12a. Mesonotum thinly tomentose in center, with erect 
hairs distributed evenly over disc and subequal in 
length to antennal segments 1 and 2 together; lat- 
eral bristles of mesonotum mostly black; males with 
abdomen mostly tomentose, each tergite with dark 
grayish-brown tomentum anteriorly and light brown- 
ish-gray fasciae posteriorly __ thologaster new species 

Mesonotum bare of tomentum in center, with erect 
hairs mostly confined to dorsocentral rows and sub- 
equal in length to antennal segment 1; lateral bristles 
of mesonotum all white to brownish yellow; males 
with abdominal tergites mostly bare in center and 
tomentose posterolaterally ____ janiceae new species 


12b. 


DESCRIPTIONS OF TAXA.—The genus Zabrops 
is readily divisible into three distinctive species 
groups which are probably equivalent to subgen- 
era. However, to treat them as formal subgenera 
would serve little purpose in a genus with only 
five known species. 


FLAVIPILIS SPECIES GROUP 


This group, containing the single species— 
flavipilis (Jones)—is characterized by the follow- 
ing features. 

Head: face narrow, eye 1.5 to 1.7 times wider; 
lower 55 to 60 percent of face gibbous; antennae 
long, 90 percent height of eye, segment 3 narrow, 
three times as long as wide; style long, with two 
segments (the first very short). 

Legs: hind tibiae with ventral surface thickly 
covered with soft pile of uniform height. 

Wings: first posterior cell closed at margin; 
costa complete. 

Abdomen: tergites 1 and 2 with lateral bristles 
(rather fine on tergite 2). 

Male genitalia: hypandrium very large, three- 
fourths as wide as gonopods; hypandrial lobes 
wide, subcontiguous, and emarginate apically; 
gonopods with median process bifid, each half 
widely separated at base and converging or cros- 
sing apically; lateral processes with large pos- 
terolateral flange on club; posterolateral mar- 
gins of epandrium produced into long points; 
aedeagus with tubes straight on distal half. 


FISHER: NORTH AMERICAN LAPHYSTIINI 


44 


49 


HK. 1.0 mm ————_—_ 


Figures 43 to 45. Male genitalia of Zabrops flavipilis. Fig. 43. Gonopods and hypandrium, ventral view, median process 
stippled, hypandrium lined. Fig. 44. Epandrium, dorsal view. Fig. 45. Aedeagus, lateral view, ventral surface above. 


Zabrops flavipilis (Jones) 

(FIGURES 43 to 45, 58, 74) 

Triclis flavipilis Jones, 1907: 275. [Type locality: Meadow, 
Nebraska. Holotype: ?University of Nebraska (not 
examined). ] 

Triclis flavipilis, Back, 1909: 232; Hull, 1962: 86. 

Zabrops flavipilis, Martin and Wilcox, 1965: 386. 


DIAGNosis.—A_ shiny-black species with 
sparse golden-yellow pile and tomentum. The 
wide hypandrium, with the lobes apically emar- 
ginate, and the shape of the gonopods are diag- 
nostic. 

MALE.—Head black; face densely golden to- 
mentose, frons, vertex, and occiput more thinly 
so, lower half of frons with transverse shiny 
band; mystax black on upper third, remainder 
golden-yellow; hairs golden-yellow on plane por- 
tion of face, ocellar tubercule, vertex, and oc- 
ciput; bristly hairs on sides of frons and slender, 
somewhat proclinate, occipital bristles black; 
beard and hairs on proboscis and palpi yellowish 
white. Antennal segment | ventrally with many 
long black bristles and a few yellow hairs, dor- 
sally with a few short yellow hairs; apex of seg- 
ment 2 witha few short black hairs; length ratio of 
segments 1.5:1.0:2.5:0.5. 


Thorax black, humeri and postalar calli brown- 
ish; dorsum shiny, golden-yellow tomentum 
present on thin lateral margin of pronotum and 
mesonotum and on two small spots above scutel- 
lum; pleura with tomentum yellowish-gray; lat- 
eral mesonotal bristles black, remainder of 
thoracic pile and bristles golden-yellow; pile 
short, sparse, and recumbent anteriorly on meso- 
notum, longer posteriorly and on pleura; two 
to three short anterior dorsocentrals. Scutellum 
with a few short hairs on disc and 12 long, slen- 
der, marginal bristles. Legs black, pile golden- 
yellow, bristles mostly same but with some black 
on tibiae and tarsi; ventral surface of pro- and 
mesotibiae with scattered pile and long bristles, 
of metatibiae with dense brush of short pile. 
Wings subhyaline, anterior margin and apical 
one-third slightly infuscate; veins dark brown. 

Abdomen (Fig. 58): black, thin apical margins 
of tergites 4 to 6 brownish; sides of tergite 1 and 
posterolateral corners of tergites 2 to 5 with 
golden-yellow tomentose spots, these spots be- 
coming shorter and wider posteriorly; sternites 
mostly shiny, each with posterior margin 
yellow-gray tomentose; pile golden-yellow, short 
and very sparse dorsally, longer laterally and 


200 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


ventrally. Genitalia: as in Figs. 43 to 45; basistyli 
and epandrium with bristles black. 
FEMALE.—No specimens examined. Judging 
from the original description, the female differs in 
having the mystax mostly black, with some yel- 
low bristles near the oral margin. In addition, 
probably the face and abdomen are wider and the 
tomentose markings are more extensive on the 
abdominal tergites; these differences occur in the 
females of the other species of Zabrops. 


SPECIMENS EXAMINED (Two males) KANSAS, Riley 
County: 13 July (Popenoe, EF). OHIO, Scioto County: Friend- 
ship, 16 July 1961 (P. H. Freytag, OSU). 


Discussion.—I have not examined the female 
holotype but Jones’s (1907) excellent description 
leaves no doubt as to the identity of this distinc- 
tive species. 

Other than differences in size, there is only 
slight variation between the two males studied. 
The Ohio specimen has more deeply emarginate 
hypandrial lobes than shown in Fig. 43 (Kansas 
male) and the median processes of the gonopods 
cross apically. The Kansas specimen has red 
humeri and postalar calli, and the two tomentose 
spots above the scutellum are much larger. 

Measurements (in mm; values without pa- 
rentheses of Kansas specimen, those with pa- 
rentheses of Ohio specimen): eye width 1.00 
(0.88); face width 0.60 (0.56); body length 10.1 
(9.2); wing length 5.8 (6.0); tergite 3 width 2.1 
(1.9). Ratios: eye width/face width 1.67 (1.57); 
body length/wing length 1.75 (1.53). 

Zabrops flavipilis is by far the most widely 
distributed species in the genus, the three known 
specimens indicating a range of at least 1400 km 
across the central United States (Fig. 74). In fact, 
the spotty distribution of this species, with its 
known range far removed from other Zabrops, 
seems to indicate that it is a relict species. It also 
appears to be the most primitive species in the 
genus, as shown by these characteristics: the 
long, two-segmented antennal style; the com- 
plete costa; the large, wide hypandrium, which is 
scarcely reduced or emarginate. 


TAGAX SPECIES GROUP 


A group consisting of two polytypic species— 
tagax (Williston) and wilcoxi new species—and 
characterized as follows. 

Head: eye about 1.1 to 1.4 times wider than 
face; lower 65 to 70 percent of face gibbose; an- 
tennae shorter, about two-thirds height of eye, 
segment 3 wider, 2.5 times longer than wide; style 
short, with only one segment. 

Legs: ventral surface of hind tibiae thickly 
covered with soft pile of uniform height. 

Wings: first posterior cell closed and usually 
short petiolate; costa complete. 

Abdomen: tergites | and 2 or | to 3 with strong 
lateral bristles. 

Male genitalia: hypandrium small, less than 
half as wide as gonopods; hypandrial lobes nar- 
row, about 1.8 to 2.8 times longer than wide, 
widely separated and rounded apically; gono- 
pods with median process entire or with apex 
notched; lateral processes with club small, with- 
out flange; epandrial margin not produced; 
aedeagus with tubes curved on distal half. 


Zabrops tagax (Williston) 


A partly shiny-black species with somewhat 
dense, short, mostly recumbent, varicolored 
pile and white to yellow tomentum. Males have 
gonopods with a median process that is wider 
than long—about half as long as the lobes of the 
hypandrium—and with a deep apical notch; each 
hypandrial lobe is about 1.8 times longer than 
wide. 

This species comprises two subspecies: Z. f. 
tagax is a Mojave and Sonoran desert form; Z. f. 
argutus, new subspecies, is cismontane, occur- 
ring along the western edge of the Peninsular and 
Transverse ranges in northern Baja California 
and southern California (Fig. 73). 


Zabrops tagax tagax (Williston), new status 
(FIGURES 33 to 42, 59, 66, 73) 


Triclis tagax Williston, 1883: 9, pl. 1, figs. 6 (wing) and 6a 
(head). [Type locality: Kern County, California. Holotype: 
UK.] 


— 


Ficures 46 to 51. Male genitalia of Zabrops spp., tagax group. Fig. 46. Gonopods and hypandrium of Zabrops tagax argutus, 
ventral view, median process stippled, hypandrium lined. Fig. 47. Gonopods and hypandrium of Zabrops wilcoxi wilcoxi, ventral 
view, median process stippled, hypandrium lined. Fig. 48. Epandrium of Zabrops wilcoxi wilcoxi, dorsal view. Fig. 49. Aedeagus of 
Zabrops wilcoxi wilcoxi, lateral view, ventral surface above. Fig. 50. Gonopods and hypandrium of Zabrops wilcoxi playalis, 
ventral view, median process stippled, hypandrium lined. Fig. 51. Gonopods and hypandrium of Zabrops wilcoxi arroyalis, ventral 
view, median process stippled, hypandrium lined. 


FISHER: NORTH AMERICAN LAPHYSTIINI 


202 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


Triclis tagax, Williston, 1886: 37; Back, 1909: 231, pl. 9, fig. 3 
(whole insect). 

Zabrops tagax, Hull, 1957: 90; Wilcox, 1960: 329; Hull, 1962: 
99, figs. 58 (antenna), 444 (wing), 883, 892 (head); Martin and 
Wilcox, 1965: 386. 


DIAGNOosIs.—The nominate subspecies is 
characterized by having generally lighter-colored 
pile, the erect mesonotal hairs sparser, and the 
tomentum more extensive dorsally on the abdo- 
men. The females have whitish tomentose bands 
across the apex of each abdominal tergite; these 
bands are greatly narrowed or, usually, inter- 
rupted in the center. 

FEMALE.—Description based on_ holotype 
(specimen teneral). Length 7.9 mm. Head dark 
brown; tomentum whitish, densest along orbits, 
thinnest on vertex, occiput, and antennal seg- 
ment 1; all hairs and bristles yellowish white, 
except on bristle on ocellar tubercule black and 
beard and hairs of proboscis and palpi white; eye 
1.2 times wider than face. Antennal segment 1 
and 2 equal in length; segment 3 missing. 

Thorax black; vestiture yellowish white, lat- 
eral bristles and some posterior dorsocentrals 
black. Mesonotum with broad median stripe and 
adjacent lateral intermediate stripes shining; re- 
maining area tomentose—anterolateral corners 
(wider and longer than humeri), broad lateral 
margins, and most of posterior declivity; pile 
fairly dense, short and recumbent; erect bristly 
hairs mostly in dorsocentral rows, anteriorly 
subequal in length to antennal segment 1. Scutel- 
lum with anterior corners tomentose, abundant 
discal pile, and 14 long marginal bristles. Legs 
dark brown, hairs and bristles mostly yellowish 
white, some tarsal and metatibial bristles black. 
Wings: length 5.4 mm; hyaline, veins yellow- 
brown; first posterior cell closed at margin on one 
wing, narrowly petiolate on other. Halteres 
yellow-brown, slightly darker at base. 

Abdomen (Fig. 66): 2.0 mm wide; segments 1 
to 3 black, 4 to 7 dark brown, all with narrow 
posterior margins yellowish brown; vestiture yel- 
lowish white. Tergite | tomentose, with a small, 
subtriangular, posteromedian bare spot; tergites 
2 to 4 with lateral and posterior margins tomen- 
tose, tomentum in form of fasciae, very short 
medially and increasing in length laterally, an- 
terior corners and broad anteromedian semicir- 
cular area bare; tergites 5 and 6 similar but fasciae 
interrupted medially; venter evenly and thinly 
tomentose; pile short and recumbent dorsally, 
longer laterally on tergites 1 and 2, and ventrally. 


MALE.—Description based on specimen from 
Yermo, San Bernardino County, California, dif- 
fering from holotype in following manner. Head 
with pile whiter; length ratio of antennal seg- 
ments. 1:221)10:2:2:0:45 

Thorax with some anterior erect bristly hairs 
black; about half of scutellar bristles black. 
Wings with veins dark brown; first posterior cells 
closed, short petiolate. 

Abdomen (Fig. 59): tergite 1 tomentose on lat- 
eral edges and narrowly along anterior margin; 
tergites 2 to 6 with tomentum confined to spots on 
posterolateral corners, these spots becoming 
shorter and wider posteriorly, except on tergite 6, 
where they are very small; on tergite 3, spots 
separated by at least half the width of abdomen. 
Genitalia (Figs. 33 to 37): median process of 
gonopods only slightly wider than long. 


SPECIMENS EXAMINED (212 males, 178 females).— 
Holotype: female, Kern County, California. 

Other specimens. USA.—CALIFORNIA, San Bernardino 
County: Yermo; Barstow; Adelanto; Goffs; (25 Mar. to 25 
Apr.). Los Angeles County: Piute Butte, (12 May). Riverside 
County: 10 miles (16 km) E of Whitewater; Indio; 5 miles (8 km) 
W of Indio; Thousand Palms; Palm Springs; (26 Feb. to 29 
Apr.). San Diego County: Borrego Valley; Coyote Creek, 4.5 
miles (7.2 km) N of Borrego Springs; Borrego Springs; Tub 
Canyon; Ocotillo; (7 Mar. to 9 Apr.). Imperial County: Truck- 
haven; (5 Apr.).—ARIZONA, Yuma County: Ehrenberg; 10 
miles (16 km) W of Aguila; Wellton; (6-15 Apr.). Pinal County: 
1.5 miles (2.4 km) N of Florence; Florence; 5 miles (8 km) S of 
Florence; Florence Junction; (9 Apr. to 2 May). Maricopa 
County: Tempe; 8 miles (13 km) S of Buckeye; Palo Verde; 
Sentinel; 18 miles (29 km) S of Gila Bend; (7-22 Apr.). Pima 
County: Marana; 23 miles (37 km) N of Ajo; 10 miles (16 km) N 
of Ajo; Organ Pipe Cactus National Monument; (16 Mar. to 12 
Apr.). MEXICO.—Sonora: 27 miles (43 km) SE of San Luis; 
25 miles (40 km) NW of Sonoita; between Sonoita and Punta 
Penasco, 500 feet (152 m); (2-30 Mar.). 


DiscussIon.—The holotype represents a pop- 
ulation that apparently has not been rediscov- 
ered. The only locality information accompany- 
ing the type is ‘‘Kern Co Clf.’’ No additional 
specimens of Z. t. tagax have been collected 
in Kern County. Also, in the holotype the to- 
mentose abdominal fasciae is entire, and the 
mystax, anterior mesonotal pile and scutellar 
bristles are all yellowish white. Several female 
specimens collected in the Mojave Desert (at 
Yermo and Adelanto) approach this phenotype; 
however, they have narrowly interrupted ab- 
dominal fasciae. All other populations of this 
subspecies studied have some of the above 
mentioned hairs or bristles partly black. 

Variation in populations of Z. t. tagax is as 


FISHER: NORTH AMERICAN LAPHYSTIINI 203 


LK 


y 


|———————_ 1.0 mm 


04 


et 
Yi 


yt 
Vi 
we 


96 57 


FiGurEs 52 to 57. Male genitalia of Zabrops spp., thologaster group. Fig. 52. Gonopods and hypandrium of Zabrops thologaster, 
ventral view, median process stippled, hypandrium lined. Fig. 53. Epandrium of Zabrops thologaster, dorsal view. Fig. 54. 
Aedeagus of Zabrops thologaster, lateral view, ventral surface above. Fig. 55. Aedeagus of Zabrops janiceae, lateral view, ventral 
surface above. Fig. 56. Epandrium of Zabrops janiceae, dorsal view. Fig. 57. Gonopods and hypandrium of Zabrops janiceae, 
ventral view, median process stippled, hypandrium lined. 


204 


follows: CALIFORNIA: San Bernardino County: 
males with scutellar bristles sometimes all black; 
posterolateral spots frequently smaller, sepa- 
rated by three-fourths the width of abdomen on 
tergite 3. Females with mystax all white to half 
white, half black; scutellum with many to all bris- 
tles yellow; fasciae on abdominal tergites 2 to 4 
usually interrupted by about one-fourth the width 
of abdomen. Riverside County, San Diego 
County: males with scutellar bristles sometimes 
all black; posterolateral spots on tergites 2 to 5 
more approximate medially, separated by about 
one-third the width of abdomen. Females gener- 
ally with mystax mostly, and scutellar bristles 
entirely, black; abdominal fasciae entire or 
slightly interrupted. ARIZONA: males with scutel- 
lar bristles frequently all black; abdominal spots 
usually separated by half the width of abdomen 
on tergite 3. Females with mystax one-third to 
one-half black; abdominal fasciae interrupted by 
one-fourth the width of abdomen; occasionally 
(two specimens from Pinal County) anterior por- 
tion of tergites 2 to 4 thinly tomentose, with 
middle one-fourth bare. SONORA: males with 
many to all scutellar bristles yellow; abdominal 
spots separated on tergite 3 by two-thirds to 
three-fourths the width of abdomen. Females 
with mystax all white; scutellar bristles all or 
nearly all yellow; tergite 1 with posteromedian 
bare area twice as broad as in holotype, and 
tergites 2 to 5 with fasciae separated by one- 
third the width of abdomen. 

The female specimens from Sonora are simi- 
lar to females of the new subspecies described 
below, at least in respect to the tomentose pat- 
terns of the abdomen. 


Zabrops tagax argutus Fisher, new subspecies 
(FiGureEs 46, 60, 67, 73) 


DIAGNOs!Is.—Distinguished from Z. t. tagax 
by the darker pile, the denser erect hairs of the 
mesonotum, and the reduced tomentum on the 
abdominal tergites of the females. 

HoLtotyre MALE.—Length 10.4 mm. Differs 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


from Z. ¢. tagax in the following respects. Light 
colored vestiture of body generally with more 
yellow caste; erect bristly hairs of mesonotum 
black, dense, equally distributed over disc, an- 
teriorly subequal in length to antennal segments | 
and 2 together. Scutellum with marginal bristles 
black. Abdomen (Fig. 60) with sides only of ter- 
gite | tomentose; tergites 1 to 6 with some black 
pile centrally. Genitalia as in Figure 46; median 
process almost twice as wide as long. 

ALLOTYPE FEMALE.—Length 9.7 mm. Similar 
to male. Head with these hairs and bristles black: 
mystax (except for a few hairs laterally and on 
oral margin yellow); several strong bristles on 
antennal segments | and 2; most of hairs on side 
of frons; bristles on ocellar tubercule. Abdomen 
(Fig. 67) with tergite | tomentose laterally and 
anteriorly, leaving a large triangular bare area; 
tergites 2 to 6 with tomentum broadly inter- 
rupted, at tergite 3 by about one-third the width of 
abdomen. 

TYPE SPECIMENS.—Holotype (CAS): male, 7 miles (11 km) 
NW of El Rosario, Baja California, Mexico, 28 Mar. 1970 (E. 
M. and J. L. Fisher, EF). Allotype female (CAS), same data as 
for holotype. 

Paratypes (32 males, 13 females). MEXICO.—BajJa CALI- 
FORNIA: 9 males, 2 females, same data as for holotype; 10 miles 
(16 km) NW of El Rosario, 1 male, same date and collector as 
for holotype; near Consuelo, 6 miles (9.7 km) NW of El Rosa- 
rio, 2 males, 3 females, 18 Apr. 1965 (D. Q. Cavagnaro, C. F. 
and E. S. Ross, and V. L. Vesterby, CAS); 1 mile (1.6 km) E of 
El Rosario, 1 male, 9 Apr. 1969 (S. C. Williams, CAS); 18.3 km 
E of El Rosario, 1 male, 9 Apr. 1973 (E. L. Sleeper, CSLB); La 
Bocana Beach, 7 km W of El Rosario, | female, 12 Apr. 1973(E. 
L. Sleeper, CSLB). USA.—CALIFORNIA, Riverside County: 
Lake Mathews, | male, 16 Apr. 1953 (A. L. Melander, USNM); 
8 miles (12.9 km) E of Sunnymead, 2 males, 12 Apr. 1958 (E. I. 
Schlinger, UCD, CM); Riverside, 3 males, 3 females, 12 May 
1949 (A. L. Melander, USNM). Los Angeles County: Azusa, | 
male, 1 female, 24 May 1925 (collector unknown), 4 males, 2 
females, 9 May 1941 (J. Wilcox, JW); Beverly Glen Canyon, 
Santa Monica Mountains, 7 males, 4 May 1962 (E. M. Fisher, 
EF); Topanga Canyon, 1 male, 10 June 1952 (R. Laxineta, 
LACM). 


Discussion.—Variation is slight in this sub- 
species. Some of the males from the type locality 
have black hairs laterally on the frons and dor- 
sally on antennal segments | and 2. In males from 


= 


FIGURES 58 to 72. Abdominal tergites 1 (upper) and 3 (lower) of Zabrops spp., dorsal view, tomentose areas stippled (Figs. 58 to 
65 males, Figs. 66 to 72 females). Fig. 58. Zabrops flavipilis. Fig. 59. Zabrops tagax tagax. Fig. 60. Zabrops tagax argutus. Fig. 61. 
Zabrops wilcoxi wilcoxi. Fig. 62. Zabrops wilcoxi playalis. Fig. 63. Zabrops wilcoxi arroyalis. Fig. 64. Zabrops thologaster. Fig. 65. 
Zabrops janiceae. Fig. 66. Zabrops tagax tagax. Fig. 67. Zabrops tagax argutus. Fig. 68. Zabrops wilcoxi wilcoxi. Fig. 69. Zabrops 
wilcoxi playalis. Fig. 70. Zabrops wilcoxi arroyalis. Fig. 71. Zabrops thologaster. Fig. 72. Zabrops janiceae. 


4 
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206 PROCEEDINGS OF THE CALIFORNIA, ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


Los Angeles and Riverside counties the apical 
abdominal tergites tend to be reddish posteriorly, 
and some of the vestiture is more brownish yel- 
low, especially the tomentum of the mesonotum. 
Females from these counties have antennae and 
frons all yellow haired, and in one specimen from 
Riverside the apical half of the abdomen is 
broadly red. 

The subspecific name is derived from Latin 
and means “‘shiny”’ or ‘‘clear’’—in reference to 
the more shiny aspect of the abdominal tergites. 

The specimens from 7 miles (11 km) northwest 
of El Rosario were found resting on very sandy 
soil between large shrubs in the bottom of a 
canyon. Those from Beverly Glen Canyon were 
taken resting on the clay soil of a road cut. 


Zabrops wilcoxi Fisher, new species 


A species closely related to Z. tagax, and gen- 
erally quite similar in appearance, differing main- 
ly in the form of the male genitalia. In males the 
median process of the gonopods is more elon- 
gate, twice as long as wide, subequal in length to 
the lobes of the hypandrium, and the apex is 
rounded or truncate; the hypandrial lobes are 
narrower, approximately 2.6 to 2.8 times longer 
than wide. 

None of the females have an abdomen as ex- 
tensively tomentose as in Z. t. tagax; however, 
their patterns of tomentum are very similar to 
those in Z. ¢. argutus. 

Zabrops wilcoxi comprises three subspecies. 


Zabrops wilcoxi wilcoxi Fisher, new species 
(FIGURES 47 to 49, 61, 68, 73) 


DIAGNosiIs.—Distinguished from related sub- 
species by both sexes having entirely yellow 
mystax and predominantly black apical abdomi- 
nal tergites; in males the median process of the 
gonopods is rounded apically and the hypandrial 
lobes are parallel. 

HoLotyPe MALE.—Length 10.0 mm. Integu- 
ment black, narrow apices of abdominal tergites 4 
and 5, and most of 6, brown. Head with pile and 
tomentum whitish yellow; mystax dense, with 
about 60 bristles. 

Thorax with tomentum brownish yellow; re- 
cumbent and lateral pile whitish yellow; erect 
bristly hairs mostly black, fairly dense and evenly 
distributed over mesonotum, anteriorly a little 
longer than antennal segment 1. Scutellar bristles 
mostly black. Legs with pile mostly yellow; 
metatibiae with one-third of pile and two-thirds 


of bristles black; bristles of hind tarsi mostly 
black. 

Abdomen (Fig. 61) with sides of tergite 1 and 
posterolateral corners of tergites 2 to 5 whitish 
tomentose; tergite 6 bare of tomentum. Dorsal 
pile recumbent, fairly dense, about two-thirds as 
long as antennal segment |; small central area on 
each tergite black pilose, remainder yellowish- 
white pilose. Genitalia as in Figs. 47 through 49; 
apex of median process rounded; hypandrial 
lobes parallel. Strong bristles black. 

ALLOTYPE FEMALE.—Length 9.2 mm. Differs 
from male as follows: integument brown on pos- 
terior half of tergite 6; thorax with anterior 
erect bristly hairs mostly yellow, shorter, rather 
sparse; abdomen (Fig. 68) with tomentose fasciae 
interrupted on tergite 3 by about half the width. 


Type SPECIMENS.—Holotype (CAS): male, Mount Diablo, 
Contra Costa County, California, 2 June 1940 (M. Cazier, 
CAS). Allotype female (CAS), same data as for holotype. 

Paratypes (98 males, 56 females), all from California. Contra 
Costa County: 10 males, 7 females, same data as for holotype; 
Mount Diablo, 1 male, 21 May 1937 (E. S. Ross, CAS), 1 male, 
29 Apr. 1917 (E. P. Van Duzee, CAS), 1 male, 17 May 1958 (J. 
A. Chemsak, CIS); Marsh Canyon, E. of Mount Diablo, 1 
female, 20 Apr. 1947 (C. W. Anderson, CIS); near Murietta 
Caves, Mount Diablo, 8 males, 3 females, 2 June 1940 (B. 
Brookman, CAS); 3 miles (4.8 km) S of Byron, | male, 5 Apr. 
1964 (R. W. Thorpe, CIS); Danville, 4 males, 3 females, 27 Apr. 
1949, 4 males, 3 females, 22 May/5 June 1952 (F. X. Williams, 
CAS); 2 miles (3.2 km) SW of Martinez, 1 male, 1 female, 7 May 
1959 (T. R. Haig, CDA). Glenn County: Fruto, 2 males, 6 June 
1954(M. Wasbauer, CIS). Mendocino County: 10 miles (16 km) 
E of Capella, 1 female, 5 May 1960 (T. R. Haig, CDA). Lake 
County: Clear Lake Oaks, 1 male, 10 May 1959 (C. H. Laton, 
UCD). Napa County: Samuel Springs, 1 male, 9 May 1955 (R. 
M. Bohart, UCD), 2 males, 2 females, 22 May 1956 (E. I. 
Schlinger, UCD), 2 males, 2 females, 30 May 1955 (R. C. 
Bechtel, A. A. Grigarick, UCD, CM). Yolo County: Davis, 1 
male, 10 May 1960 (F. D. Parker, UCD), 1 male, 11 May 1954 
(E. I. Schlinger, CM), 1 male, 20 May 1952 (H. F. Robinson, 
UCD). Sacramento County: North Sacramento, | male, 19 
May 1955(P. D. Hurd, CIS). Solano County: Vallejo, 1 male, 27 
May 1950 (J. N. Simons, CIS). Alameda County: Berkeley, | 
female, May 1940 (J. Hansen, CAS). San Joaquin County: 
Tracy, 1 male, 15 May 1933 (A. Michelbacher, CIS). Stanislaus 
County: 3 miles (4.8 km) NW of La Grange, | male, 6 Apr. 1966 
(S. M. Fullerton, EF). San Mateo County: Redwood City, 2 
males, 2 females, 2 June 1951, 8 males, 30 May 1954 (P. H. 
Arnaud, Jr., CAS, JW). Santa Clara County: Stanford Univer- 
sity, 1 male, 3 May 1956, 2 males, 2 females (‘‘resting on 
ground, grassy hillside’), 6 May 1954, 4 males, 1 female, 18 
May 1951 (P. H. Arnaud, Jr., CAS, JW), 2 males, 1 female, 7/14 
May 1910 (W. M. Mann, USNM), 7 males, 5 females, 19/20 
May 1920, 9 males, 5 females, June 1920(F. R. Cole, CIS, JW); 
Palo Alto, 3 males, 1 female, 19 May 1940 (E. S. Ross, CAS); 
Alum Rock Park, 1 male, 25 May 1950 (J. W. MacSwain, CIS); 
San Jose, 1 female, 18 May 1965 (R. P. Allen, CIS). Merced 
County: Panoche Hills, 3 males, 3 females, 30 Apr. 1922 (E. C. 
Van Dyke, CAS). San Benito County: Panoche Creek, 3 males, 


FISHER: NORTH AMERICAN LAPHYSTIINI 


3 females, 30 Apr. 1922 (A. J. Basinger, CAS). Monterey 
County: Salinas, 5 males, 5 females, 25 May 1952 (P. H. Ar- 
naud, Jr., JW); Bryson, 1 male, 1 female, 19 May 1920 (E. P. 
Van Duzee, CAS). Fresno County: Mercy Hot Springs, | male, 
12 May 1954 (O. Bryant, CAS). Tulare County: Porterville, 1 
male, 2 Apr. 1960 (E. Ball, FSCA). San Luis Obispo County: 
Pozo, | female, 1 May 1962 (P. D. Hurd, CIS). 


DiIscUSSION.—Zabrops w. wilcoxi is found in 
the vicinity of San Francisco and San Pablo bays 
and at low elevations in the Coast Ranges to the 
north and south. Two collections of this subspe- 
cies are known from the northern part of the San 
Joaquin Valley and two specimens have been col- 
lected on the east side of the valley, near La 
Grange and at Porterville (Fig. 73). 

Several female specimens from the type lo- 
cality have all yellow scutellar and hind tibial 
bristles. In specimens of both sexes from Napa 
and Glenn counties, especially males, the apical 
one or two abdominal tergites are mostly reddish 
brown. In specimens from San Mateo and Santa 
Clara counties southward the anterior mesonotal 
hairs, the scutellar bristles, and the metatibial 
pile and bristles are all, or nearly all, yellow. 

It is a great pleasure to name this species in 
honor of Mr. Joseph Wilcox, who has contrib- 
uted so much to the knowledge of North Ameri- 
can Asilidae. 


Zabrops wilcoxi playalis Fisher, new subspecies 


(FIGURES 50, 62, 69, 73) 


D1AGNosis.—The absence of tomentum on the 
lateral portions of the humeri in both sexes and 
males with the upper one-third of the mystax 
black are characters which serve to separate this 
subspecies from others in the tagax group. 

HoLotyPeE MALE.—Length 9.4 mm. Integu- 
ment black, except tarsi brown and posterior 
one-third of tergite 4, half of 5, and all of 6 red. 
Head: tomentum grayish white; upper one-third 
of mystax, most of hairs on front and bristles on 
ocellar tubercule black; occipital bristles yellow; 
beard white; other pile yellowish white. 

Thorax with tomentum gray dorsally and yel- 
low laterally; lateral portion of humeri and area 
posterior to humeri bare of tomentum, shiny 
black; recumbent and lateral pile yellow; erect 
bristly hairs dense, black, subequal in length to 
antennal segment 1. Scutellar bristles black. 
Legs: posterior pair with bristles and short pile 
black on apical part of femora and most of tibiae; 
posterior tarsi with pile and bristles black, an- 


207 


terior and middle tarsi with some bristles black; 
remaining vestiture whitish yellow. 

Abdomen (Fig. 62): tomentum white; dorsal 
recumbent pile about as long as antennal segment 
1, black medially, yellowish on anterior and lat- 
eral margins; longer pile and bristles on tergites | 
and 2 whitish. Genitalia as in Fig. 50; median 
process tapered and somewhat truncate apically; 
hypandrial lobes slightly divergent; numerous 
strong bristles of gonopods and epandrium black. 

ALLOTYPE FEMALE.—Length 9.3 mm. Similar 
to male but differing as follows: face and front 
with whitish-yellow vestiture, mystax nearly all 
black; hind legs with short pile mostly yellow; 
abdomen with tomentum more extensive, as in 
Fig. 69; fasciae on tergite 3 separated by half 
width of abdomen. 


Type SPECIMENS.—Holotype (CAS): male, Oceano, San 
Luis Obispo County, California, 23 June 1970 (J. Wilcox, JW). 
Allotype female (CAS), same data as for holotype. 

Paratypes (138 males, 91 females), all from California. San 
Luis Obispo County: 7 males, 3 females, same data as for 
holotype; type locality, 5 males, 1 female, 9 July 1969, 4 males, 
4 females, 14/16 July 1970 (J. Wilcox, JW), 2 males, 5 females, 
24 Apr. 1951 (R. M. Bohart, UCD); Morro Dunes, 3 males, 7 
females, 17 June 1947(A. L. Melander, USNM); | mile (1.6 km) 
N of Morro Bay, | male, 1 female, 23 June 1970 (J. Wilcox, JW); 
Morro Bay, | male, 30 Apr. 1962 (J. Powell, CIS); 4 males, 2 
females, 26 June 1963, 3 males, 4 females, 4 July 1963 (J. C. 
Hall, UCR); Baywood Park, | male, 23 June 1970, 3 males, 8 
July 1969, 1 male, 1 female, 15 July 1970, 2 males, 20 Sept. 1968 
(J. Wilcox, JW); Los Osos, 1 female, 26 May 1970, 1 male, 21 
Sept. 1968 (J. Wilcox, JW); Pismo Beach, 29 males, 5 females, 
13 May 1956 (J. Wilcox, JW), 10 males, 7 females, 29 June 1959 
(P. H. Arnaud, Jr., CIS, CDA), 9 males, 4 females, 7/9 July 1969 
(J. Wilcox, JW), 1 male, 1 female, 13 July 1965 (C. A. Toschi, 
M. J. Tauber, CIS), 1 female, 15 July 1967 (M. E. Irwin, UCR), 
1 female, 26 July (J. Wilcox, JW); Grover City (‘dunes associa- 
tion’’), 35 males, 20 females, 4 July 1956, 3 males, 3 females, 14 
Aug. 1957 (P. D. Hurd, E. G. Linsley, CIS); Oso Flaco Lake, 5 
miles (8 km) S of Oceano, 4 males, 27 Apr. 1968 (J. Chemsak, J. 
Powell, CIS), 3 males, 5 females, 11 May 1965 (R. Langston, J. 
Powell, CIS), 2 males, 29 June 1967 (M. E. Irwin, UCR), 1 
male, 2 females, 13 July 1959(R. M. Bohart, F. D. Parker, CM), 
2 females, 14 July 1965 (M. R. Gardner, J. Powell, CIS, UCD); 
Dunes Lake, 7 miles (11.3 km) S of Oceano, 4 males, 6 females, 
4/5 June 1971 (J. D. Pinto, UCR). Santa Barbara County: Point 
Conception, | male, 1 female, 20 June 1951 (K. W. Tucker, 
UCD). 


Discussion.—Zabrops w. playalis inhabits 
the coastal sand dunes of Morro and San Luis 
Obispo bays. It also has been collected at Point 
Conception, so presumably it will be found in 
other suitable areas of the central California 
coast (Fig. 73). Its subspecific name is derived 
from the word ‘‘playa,’’ which is ‘‘shore’’ or 
‘‘beach’’ in Spanish. 


208 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


1200 kms 


800 miles 


w. wilcoxi 
w. Playalis 
Ww. arroyalis 
t. tagax 

t. argutus 


thologaster 


o> 8 © + Oo 


jJaniceae 


6) Miles 


0 Kilometers 


Ficures 73 and 74. Fig. 73. Distribution of Zabrops wilcoxi, tagax, thologaster and janiceae in the southwestern United States 
and northwestern Mexico. Fig. 74. Distribution of Zabrops flavipilis in the central United States. 


FISHER: NORTH AMERICAN LAPHYSTINI 


Specimens of this subspecies present a very 
uniform habitus, differing noticeably only in the 
body size and the amount of red at the apex of the 
abdomen. This character varies from the apical 
three segments all red to only tergite 6 red; males 
tend to have more red than do females. 

One male specimen (Pismo Beach, 24 June 
1959) had a small bombyliid as prey. 


Zabrops wilcoxi arroyalis Fisher, new subspecies 
(Ficures 51, 63, 70, 73) 


DIAGNosIs.—This taxon can be identified by 
the following combination of characters: both 
sexes with apical two or three abdominal tergites 
red and mesonotum with lateral margin of tomen- 
tum complete; males with mystax all white and 
median process of gonopods truncate apically; 
females with mystax mostly black and fasciae on 
tergite 3 separated by one-third width of abdo- 
men. 

HoLotyrPe MALE.—Length 10.0 mm. Integu- 
ment black, dorsum of abdominal tergite 4, and 
all of 5 and 6 red. Head with vestiture white, 
occipital bristles yellow, those of ocellar tuber- 
cule black. 

Thorax with dorsal tomentum and recumbent 
pile yellowish white; lateral vestiture white; erect 
bristly hairs black, not dense but distributed 
evenly over disc, short, anteriorly about two- 
thirds as long as antennal segment 1; scutellum 
with marginal bristles all black. Legs with pile 
white, bristles of hind legs nearly all black, re- 
maining bristles white. 

Abdomen (Fig. 63) with tomentum white, 
posterolateral spots a little wider than in Z. w. 
wilcoxi, separated by about one-half width of ab- 
domen on tergite 3, recumbent dorsal pile about 
one-half as long as antennal segment 1, nearly all 
black; longer lateral and ventral pile white. 
Genitalia as in Fig. 51; median process short, 
very slightly notched at apex; hypandrial lobes a 
little more divergent than in Z. w. playalis. 

ALLOTYPE FEMALE.—Length 10.8 mm. Head: 
mystax with about two-thirds of bristles black, 
remainder yellowish white; less dense than in 
other subspecies, consisting of around 40 bris- 
tles. Vestiture of front, vertex and occiput yel- 
lowish white; remainder as in male. 

Thorax with recumbent pile denser and erect 
bristly hairs sparser than in male. 

Abdomen (Fig. 70) with tomentum more ex- 
tensive than in other subspecies of wilcoxi; fas- 


209 


ciae on tergite 3 interrupted by one-third width of 
abdomen; dorsal recumbent pile very short, 
mostly yellowish white; lateral pile reduced, con- 
sisting mostly of bristles on tergites 1 and 2. 

TYPE SPECIMENS.—Holotype (CIS): male, 10 miles (16 km) 
W of Simmler, San Luis Obispo County, California, 5 May 1962 
(R. W. Thorp, CIS). Allotype (CIS): female, same data as for 
holotype. 

Paratypes (25 males, 18 females), all from California. San 
Luis Obispo County: 16 males, 13 females, same data as for 
holotype. Fresno County: Coalinga, 9 males, 4 females, 1 May 
1941 (J. Wilcox, JW). Kern County: 18 miles (29 km) N of 
McKittrick, 1 female, 1 May 1941 (J. Wilcox, JW). 

OTHER SPECIMENS (2 females).—CALIFORNIA, Monterey 
County: | mile (1.6 km) S of Soledad, 1 female, 11 Apr. 1961 (R. 
W. Thorp, CIS); Pleyto, 1 female, 22 May 1920 (E. P. Van 
Duzee, CAS). 


DISCUSSION.—This subspecies occurs along 
the southwestern edge of the San Joaquin Valley 
and in the foothills of the adjacent Coast Ranges. 
In the western and northern parts of its range, it 
overlaps with the nominate subspecies Z. w. wil- 
coxi, although neither taxa have been collected at 
the same locality (Fig. 73). 

Variation is as follows. Several males and 
females from the type locality and one male from 
Coalinga have some black hairs on the front, 
antennae, and anterior part of the humeri. In a 
few of these, some or all of the occipital bristles 
are also black; these bristles are white to yel- 
lowish brown in all other species of Zabrops, 
save for flavipilis. In females from the type lo- 
cality, tergites 4 and 5 may be partly blackish 
dorsally. 

The two female specimens from Monterey 
County agree in most respects with the type ma- 
terial; however, they differ in having a denser 
mystax (about 60 bristles) and yellowish tomen- 
tum of the face and front, as in the nominate 
subspecies. For these reasons they have not been 
included as paratypes. 

This subspecies is named for the habitat— 
arroyos—in which nearly all of the specimens 
have been collected. 

Two specimens of Zabrops wilcoxi (1 male, 1 
female), from an uncertain locality, are probable 
intergrades between the subspecies wilcoxi and 
playalis. These specimens have the mystax and 
abdominal tergite coloration characteristic of 
playalis, but the mesonotal tomentum and the 
color of the hind tibial pile are as in wilcoxi. The 
male genitalia are also intermediate in structure, 
the median process tapering but the hypandrium 
lobes parallel. 


210 


The locality data for the two intergrades are: 
California, Santa Barbara County, 6 miles (9.7 
km) south of Pine Mountain Summit, 13 June 
1962 (L. H. Dawson, UCR). There is a Big Pine 
Mountain and a Little Pine Mountain, both in the 
San Rafael Mountains (about 32 and 22 air km 
north, respectively, of Santa Barbara), but no 
‘*Pine Mountain Summit.’> However, there is 
such a locality in Ventura County, about 5 km 
east of the Santa Barbara county line. If this is the 
actual site referred to on the label, 6 miles (9.7 
km) south (along state highway 33) would be at 
about 1200 m in the Sespe River gorge. If these 
flies are from one of the San Rafael Mountain 
sites, the elevation would be about the same. 
Whichever locality is correct, this collection rep- 
resents the southeastern-most record for Zab- 
rops wilcoxi, as well as the highest altitude that a 
specimen of the genus has been taken. 


THOLOGASTER SPECIES GROUP 


Two new species, known only from Lower 
California, form this group—Z. thologaster and 
Z. janiceae. The group can be differentiated by 
the features below. 

Head: as in the tagax group; antennae with 
segment 3 more robust, two times longer than 
wide. 

Legs: hind tibiae without soft pile. 

Wings: first posterior cell closed and short 
petiolate; costa evanescent beyond apex of wing, 
either greatly reduced in width, or entirely ab- 
sent; anterior branch of third vein (R,) sometimes 
with basal stump vein or, in rare cases, a com- 
plete crossvein forming third submarginal cell. 

Abdomen: generally tergites 1 to 4 or 1 to 5 with 
strong lateral bristles. 

Male genitalia: hypandrium moderate sized, a 
little over half as wide as gonopods; hypandrial 
lobes about 1.5 times longer than wide, well sepa- 
rated and rounded apically; gonopods with me- 
dian process bifid, the halves subcontiguous; lat- 
eral processes with club large, without flange; 
epandrium with posterolateral margins rounded, 
slightly produced; aedeagus with tubes curved on 
distal half. 


Zabrops thologaster Fisher, new species 
(FiGuREs 52 to 54, 64, 71, 73) 


DIAGNosis.—A species with extensive brown 
and gray tomentum dorsally in both sexes, the 
mesonotum covered with long bristly hairs, and 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


the males with the median process of the 
gonopods as long as the hypandrial lobes. 
HoLotyPpe MALE.—Length 7.9 mm. Integu- 
ment black, tarsi, trochanters, and narrow apices 
of tergites 3 to 5, and most of 6, brownish. Head: 
vestiture grayish white, except occipital bristles 
yellowish brown, about 20 bristles in upper third 
of mystax and two bristles of ocellar tubercule 
black. Antennae with segment | a little longer 
than segment 2, together subequal to segment 3. 
Thorax: lateral margin of mesonotum and 
pleura gray tomentose; broad median stripe and 
shorter lateral intermediate stripes thinly brown 
tomentose; these stripes separated by narrow, 
longitudinal, grayish-brown tomentose lines 
which connect to a large patch of golden-brown 
tomentum on posterior declivity. Recumbent 
dorsal pile brownish yellow, rather sparse, espe- 
cially in middle; pleural pile grayish white. Erect 
bristly hairs black, sparsely but evenly distrib- 
uted over disc, anteriorly as long as antennal 
segments | and 2 together, posteriorly as long as 
antennae. Scutellum with golden-brown tomen- 
tose spots on lateral margins, and similar colored 
discal pile; about 20 marginal bristles, about 5 
black, 15 yellowish brown. Legs: hind tibiae and 
tarsi with pile and bristles about half black, rest of 
legs with vestiture yellowish white. Wings: 5.3 
mm long, very lightly and evenly infuscated. 
Abdomen (Fig. 64): 1.7 mm wide. Tergites | to 
5 with posterior margin and most of lateral mar- 
gins gray tomentose, remainder of dorsum with 
thin grayish-brown tomentum; anterolateral cor- 
ners of tergites 2 to 5 bare; tergite 6 mostly bare, 
apical corners grayish tomentose. Short recum- 
bent pile dense, colored to match tomentose pat- 
terns; longer pile on sides of tergites | to 3 and on 
venter white. Genitalia: as in Figs. 52 to 54; bris- 
tles and hairs mixed black and white. 
ALLOTYPE FEMALE.—Length 8.2 mm. Very 
similar to male. Mystax about three-fourths 
black; mesonotum with median and _ lateral 
stripes mostly obscured by denser tomentum; 
recumbent pile denser, especially in middle; 
erect bristly hairs slightly shorter; scutellum with 
tomentose spots larger and marginal bristles all 
yellowish brown; legs with bristles nearly all yel- 
lowish white; abdomen 2.1 mm wide, with tergite 
6 tomentose as on tergites | to 5. 


Type SPECIMENS.—Holotype (CAS): male, 0.5 miles (0.8 
km) NE of El Molino, Bahia San Quintin, Baja California, 
Mexico, 29 Mar. 1970 (E. M. and J. L. Fisher, EF). Allotype 
female (CAS), same data as for holotype. 


FISHER: NORTH AMERICAN LAPHYSTINI 


Paratypes (80 males, 32 females), all from Mexico. BAJA 
CALIFORNIA: 57 males, 8 females, same data as for holotype; 3 
miles (4.8 km) NW of Camalu, 23 males, 19 females, 29 Mar. 
1970 (E. M. and J. L. Fisher, EF); El Socorro Sand Dunes, 5 
males, 12 Apr. 1973 (E. L. Sleeper, CSLB). 

OTHER SPECIMENS (all from Mexico).—BAJA CALIFORNIA: 
21 miles (34 km) SSE of El Rosarito (long. 114°W), 1 male, 9 
Oct. 1972 (E. M. Fisher, EF).—BaAJA CALIFORNIA SuR: Guer- 
rero Negro, 3 males, 18 Mar. 1967 (D. S. Verity, EF); 15 miles 
(24 km) S of San [Santo] Domingo, | male, 4 Oct. 1941 (R. M. 
Bohart, E. S. Ross, CAS). 


Discussion.—Zabrops thologaster occurs 
along the Pacific coast of Lower California where 
it has been found on very sandy soil just inland 
from the shore line (Fig. 73). Many specimens 
were taken sitting next to, or even underneath, 
low-growing plants. 

The species is the only Zabrops in which males 
have an abdomen as extensively tomentose as 
that of females. Its name is derived from the 
Greek (“‘tholos,”’ dirt or dirt-colored; and *‘gas- 
ter,’ belly) and refers to the gray-brown color of 
the abdomen. 

Considerable variation is present in_ this 
species. In specimens from the type locality, the 
mystax ranges from one-fourth to one-half black 
in males, and one-half to three-fourths black in 
females; scutellar bristles range from half black 
to all yellow in both sexes. Females from 3 miles 
(4.8 km) north of Camalu exhibit similar variation 
but may have both mystax and scutellar bristles 
mostly black. The five males from El Socorro 
Sand Dunes have all white mystax and all yellow 
scutellar bristles. 

Specimens from 21 miles (34 km) SSE of El 
Rosarito, Guerrero Negro, and 15 miles (24 km) 
south of San Domingo (all males) have all white 
mystax and all yellow scutellar bristles and these 
variations as well: mesonotum with tomentum 
thinner, partly shiny on median stripe, lateral 
stripe, and posterior declivity; abdominal tergites 
1 to 5 with a posteromedian bare spot (narrowly 
interrupting the fasciae) and tergites 2 to 5 with 
anterior margins narrowly bare; integument red- 
dish brown on ventral side of all femora and basal 
half of all tibiae. In the specimen from 15 miles (24 
km) south of San Domingo, the integument of 
abdominal segments 2 to 5 are also broadly red- 
dish apically. 

Several male and female specimens from Bahia 
San Quintin and Camalt have a small stump vein 
present at the base of the anterior branch of the 
third vein (R,). One female (Camalt) has a com- 


211 


plete crossvein forming a third submarginal cell, 
in addition to a stump vein. 

The material from the southern half of the 
range of Z. thologaster is different enough 
phenotypically to indicate that subspecific dis- 
tinction might be warranted. However, it would 
be premature to do this now as only a few indi- 
viduals (and no females) have been collected in 
this area, and the full range of variation is not 
known. Also the male genitalia of specimens 
from Guerrero Negro are the same as in the type 
series. 

Two individuals collected from Bahia San 
Quintin had small flies as prey: a male with a 
ceratopogonid and a female with a tachinid. 


Zabrops janiceae Fisher, new species 
(FicureEs 55 to 57, 65, 72, 73) 


DIAGNOosIs.—Very similar to Zabrops tholo- 
gaster, but distinguished from that species as fol- 
lows: (1) mesonotum in both sexes bare of tomen- 
tum dorsally, with erect hairs shorter and spars- 
er, and with lateral bristles all white to brownish 
yellow; (2) males with abdominal tergites exten- 
sively bare of tomentum and with median process 
of gonopods half as long as hypandrial lobes; (3) 
females with abdominal tergites light- and dark- 
gray tomentose. 

Ho.totyre MALE.—Length 9.7 mm. Integu- 
ment black; these portions red: apical one fourth 
of abdominal tergites 2 to 5 and most of 6, 
trochanters, tibiae, and tarsi. Head: face and 
front yellowish-gray tomentose; vertex and oc- 
ciput gray tomentose. Mystax and beard white; 
antennae and frontal hairs yellowish white; bris- 
tles of ocellar tubercule and occiput yellow. 

Thorax: gray tomentose, broad median dorsal 
stripe, confluent lateral-intermediate stripes, and 
narrow, median posterior stripe shining black. 
Pile white laterally, yellowish white dorsally, 
with some black median and posterior dorsocen- 
tral hairs; short, recumbent pile very dense, par- 
tially obscuring black integument, absent in very 
narrow anteromedian, longitudinal stripe; an- 
terior erect bristly hairs short—as long as anten- 
nal segment I—and very sparse. Scutellum with 
posterolateral corners white tomentose; discal 
pile and marginal bristles yellowish white. Legs: 
pile and bristles white. Wings: length 7.4 mm; 
membrane hyaline; base of R, with small stump 
vein. 

Abdomen (Fig. 65): width 1.9 mm. Lateral and 


212 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 5 


narrow anterior margins of tergite 1, and lateral 
edges and posterolateral corners of tergites 2 to 5, 
thinly gray tomentose; tergites | to 6 densely 
covered with short, recumbent, yellow pile 
which partially obscures ground color; lateral 
pile and bristles yellowish white. Genitalia: as in 
Figs. 55 to 57; bristles mostly white. 

ALLOTYPE FEMALE.—Length 9.2 mm. Very 
similar to male holotype with these differences: 
disc of scutellum with central bare spot, remain- 
der gray tomentose; abdomen (Fig. 72) entirely 
gray tomentose above, except anterolateral cor- 
ners of tergites 2 to 5 and dorsomedian oval spots 
on tergites | to 6 bare. Anterodorsal portion of all 
tergites yellowish gray, remainder whitish gray. 
The shade and intensity of tomentum color 
changes with angle of view; when viewed from 
above and without magnification, tomentum ap- 
pears dark gray anteriorly and light gray pos- 
teriorly. 

Type SPECIMENS.—Holotype (CAS): male, 20 miles (32.2 
km) W of Bahia de Los Angeles, Baja California, 22 Mar. 1970 
(E. M.and J. L. Fisher, EF). Allotype female (CAS), same data 
as for holotype. 

Paratypes (10 males, 19 females), all from Mexico. BAJA 
CALIFORNIA: 8 males, 18 females, same data as for holotype; 
Arroyo Santo Dominguito, 1.5 miles (2.4 km) S of Santa Rosalil- 


lita, 2 males, 1 female, 25 Mar. 1970 (E. M. and J. L. Fisher, 
EF). 


Discussion.—This species is known from only 
two localities in central Baja California, one on 
the Pacific coast and the other near the Gulf 
coast, east of the Peninsular Divide (Fig. 73). 

Specimens from both these localities are very 
similar, and the total variation in the material at 
hand is small. Individuals of both sexes have 
these variations: apical half of hind tibiae darker 
and with brown to black bristles; wing venation 
anomalies similar to those in Z. thologaster; lat- 
eral bristles of mesonotum yellowish brown; one 
to three extra notopleural bristles anteriorly. In 
females the bare spot on the scutellum sometimes 
extends to the posterior margin of the disc. 

The specimens from 20 miles (32.2 km) west of 
Bahia de Los Angeles were taken on sandy soil 
about ice plant, Mesembryanthemum crystal- 
linum Linneaus; those from Arroyo Santo 
Dominguito were found atop a large sand dune 
which bordered a large sandy wash, about one 
kilometer from the coast. 

Zabrops janiceae is named in honor of my 
wife, Janice Lee Fisher, who collected part of the 
type material and has collected many other inter- 
esting robber flies as well. 


LITERATURE CITED 


ArTIGAS, J. N. 1971. Las estructuras quitinizadas de la 
spermatheca y funda del pene de los asilidos y su valor 
sistematico a traves del estudio por taxonomia numerica 
(Diptera-Asilidae). Gayana Zool. (18): 1-106, figs. 1-138. 

Back, E. A. 1909. The robber-flies of America north of 
Mexico, belonging to the subfamilies Leptogastrinae and 
Dasypogoninae. Trans. Am. Entomol. Soc. (Phila.) 35: 
137-400, pls. I-XI. 

BONHAG, P. F. 1949. The thoracic mechanism of the adult 
horsefly (Diptera: Tabanidae). Cornell Univ. Agric. Exp. 
Stn. Mem. 285: 1-39, figs. 1-25. 

CRAMPTON, G. C. 1942. Guide to the insects of Connecticut. 
Part VI. The Diptera or true flies of Connecticut. Fasc. 1. The 
external morphology of the Diptera. Conn. State Geol. Nat. 
Hist. Surv. Bull. (64): 10-165, figs. 1-14. 

HERMANN, F. 1905. Beitrag zur Kenntnis der Asiliden (Dip- 
tera). Berl. Entomol. Z. 50: 14-42, figs. 1-29. 

HuLL, F.M. 1957. Some flies of the family Asilidae (Diptera). 
Psyche 64: 90-96. 

1962. Robber flies of the world. The genera of the 
family Asilidae. U.S. Nat. Mus. Bull. 224 (Pt. 1): i-x, 1-430, 
text-figs. 1-29, 1 pl.; (Pt. 2): 431-907, text-figs. 30-35, figs. 
1-2536. 

Jones, P. R. 1907. A preliminary list of the Asilidae of Ne- 
braska with description of new species. Trans. Am. En- 
tomol. Soc. (Phila.) 33: 273-286. 

Kar_, E. 1959.  Vergleichend-morphologische Unter- 
suchungen der manlichen Kopulationsorgane bei Asiliden 
(Diptera). Beitr. Entomol. 9(5/6): 619-680, figs. 1-52. 

KNuTSsoNn, L. V. 1972. Pupa of Neomochtherus angustipen- 
nis (Hine), with notes on feeding habits of robber flies and a 
review of publications on morphology of immature stages 
(Diptera: Asilidae). Proc. Biol. Soc. Wash. 85(13): 163-178, 
figs. 1-9. 

KRIVOSHEINA, N. P. 1973. Larvae and pupae of robber flies 
from the genus Laphystia (Diptera, Asilidae). Zool. Zhurn. 
52(3): 457-459, figs. 107. [In Russian.] 

Lene, P. A. 1969. Assassin flies of the tribe Laphystiini 
(Diptera, Asilidae) of the USSR fauna. Zool. Zhurn. 48(2): 
233-240, fig. 1. [In Russian.] 

Loew, H. 1847. Ueber die europaischen Raubfliegen 
(Diptera-Asilica). Linnaea Entomologica, Stettin 2: 384-568. 
[Not seen. ] 

. 1874. Neue nordamerikanische Dasypogonina. Berl. 
Entomol. Z. 18: 353-377. 

MartTIN, C. H. 1968. The new family Leptogastridae (the 
grass flies) compared with the Asilidae (robber flies) (Dip- 
tera). J. Kans. Entomol. Soc. 41(1): 70-100, figs. 1-137. 

, AND N. PAPAVERO. 1970. Family Asilidae. In A 

catalogue of the Diptera of the Americas south of the United 

States. Museu de Zoologia, Universidade de Sao Paulo 

(35B): 35B.1-35B. 139. 

, AND J. WiLcox. 1965. Asilidae. Pages 360-401 in 
Alan Stone et al., A catalog of the Diptera of America north 
of Mexico... U.S. Dept. Agric. Agric. Hdbk. (276): 360-401. 

Mayr, E. 1963. Animal species and evolution. The Belknap 
Press of Harvard University Press, Cambridge, Mass. 797 
pp. 

OLpRoyD, H. 1963. The tribes and genera of the African 
Asilidae (Diptera). Stuttg. Beitr. Naturkd. (107): I-16, figs. 
1-24. 


1970. Studies of African Asilidae (Diptera). I. 
Asilidae of the Congo Basin. Bull. Br. Mus. (Nat. Hist.), 
Entomol. 24(7): 209-334, figs. 1-96. 


FISHER: NORTH AMERICAN LAPHYSTIINI 213 


TORRE-BUENO, J.R. DELA. 1962. A glossary of entomology, 1886. Dipterological notes and descriptions. Trans. 
3rd ed. Brooklyn Entomol. Soc., Brooklyn, New York. 336 Am. Entomol. Soc. (Phila.) 13: 287-307. 
1893. New or little-known Diptera. Kans. Univ. 


pp. 
Witcox,J. 1960. Laphystia Loew in North America (Diptera: Quart. 2: 59-78. 
Asilidae). Ann. Entomol. Soc. Am. 53(3): 328-346, figs. 1-2. 1901. Supplement (part). Pages 297-328, pl. 5, figs. 
WILLISTON, S. W. 1883. On the North American Asilidae 13-14, pl. 6, figs. 1-6 in F. D. Godman and O. Salvin, eds. 
(Dasypogoninae, Laphriinae), with a new genus of Syr- Biologia Centrali-Americana, Zoology, Insecta, Diptera, 1. 
phidae. Trans. Am. Entomol. Soc. (Phila.) 11:1-35, pl. 1, London. 378 pp. 


figs. 1-15, pl. 2, figs. 1-14. 


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PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 6, pp. 215-231; 17 figs.; 1 table 


August 18, 1977 


SYSTEMATICS AND BIOLOGY OF THE DEEP-SEA FISH FAMILY 
GIBBERICHTHYIDAE, A SENIOR SYNONYM OF THE 
FAMILY KASIDOROIDAE' 


By 


Donald P. de Sylva 


University of Miami, Rosenstiel School of Marine und Atmospheric Science 
Miami, Florida 33149 


and 


William N. Eschmeyer 


California Academy of Sciences, San Francisco, California 94118 


ABSTRACT: The fish family Kasidoroidae Robins and de Sylva (1965) is shown to be based on Atlantic 
specimens of the unusual prejuvenile stage of the xenoberyciform genus Gibberichthys Parr (1933). Kasidoron 
edom Robins and de Sylva (1965) is treated as the prejuvenile stage of Gibberichthys pumilus Parr (1933). 
Kasidoron latifrons Thorp (1969), described from a prejuvenile specimen from the western Indian Ocean, is 
confirmed to be a separate species, G. latifrons; it is now known from the Indian Ocean and the western and 
southwestern parts of the Pacific Ocean. Ontogenetic changes from the larval stage to the adult are discussed 
and illustrated for G. pumilus , particularly the development and loss of the pelvic appendage—a modified third 
pelvic fin ray—and the development of crests on the head. The prejuvenile stage is found in epipelagic waters, 
and adults apparently occur in the lower mesopelagic and perhaps upper bathypelagic levels; adults probably 
do not undergo daily vertical migration. Additional notes on biology are included. 


INTRODUCTION 


A new family of fishes, the Kasidoroidae, was 
established by Robins and de Sylva (1965) for a 
small western Atlantic species characterized by a 
peculiar pelvic appendage, a modified third pel- 
vic fin ray. They included the family in the order 


' Contribution from the University of Miami, Rosenstiel 
School of Marine and Atmospheric Science. Biological Results 
of the University of Miami Deep-Sea Expeditions No. 122. 
This research was supported in part by National Science 
Foundation research grants GB-893, GB-4472, GB-8047 to the 
senior author, by research grant OCE 76-02251 to Dr. Bruce 
Robison, and by Biological Ship Time grants GB-1204 and 
GA-4569. 


Mirapinniformes. Robins (1966) provided addi- 
tional comments on the family and order. Myers 
and Freihofer (1966) compared the family 
Kasidoroidae (incorrectly spelled by them as 
Kasidoridae) with their family Megalomyc- 
teridae. Greenwood et al. (1966) listed the family 
Kasidoridae (sic) in the suborder Mirapin- 
natoidei, order Cetomimiformes. However, 
Rosen and Patterson (1969: 456) suggested that, 
‘‘from illustrations of this fish [Kasidoron edom|] 
and a brief examination of the holotype we are 
convinced that it is very closely related to, and 
may be only the larva of, the beryciform Gib- 
berichthys pumilus, with which it agrees in al- 


[215] 


216 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 6 


most every character. . . . Kasidoron is therefore 
a beryciform. . . .”’ Rosen and Patterson did not 
elaborate further. A second species of Kasido- 
ron, K. latifrons, was described by Thorp (1969) 
from off Zanzibar in the western Indian Ocean. 

During the period 1969-1972, we were able to 
examine specimens referable to Kasidoron 
edom collected since the description of that 
species and many adult specimens identifiable as 
Gibberichthys pumilus. Our study of these 
specimens confirmed that Kasidoron edom in- 
deed represented the prejuvenile stage of Gib- 
berichthys pumilus, and we were thus able to 
trace ontogenetic changes and to provide infor- 
mation on biology and distribution. Our paper 
was ready to publish in 1972 and was cited as in 
press by Ebeling and Weed (1973: 412, footnote) 
in their treatment of the family Gibberichthyidae 
in the series titled *‘Fishes of the western North 
Atlantic.’ One dilemma caused us to withhold 
the paper at that time—this involved specimens 
from areas other than the Atlantic. At that time 
(1972) we had an adult specimen from the western 
Indian Ocean and a small postlarval specimen 
from the Molucca Sea. The adult was not in good 
condition and the description of latifrons was 
based on a prejuvenile which seemed to be 
somewhat different from the Atlantic pre- 
juveniles available to us. Was latifrons a separate 
species? 

In 1973 Dr. John Paxton made available to us 
an adult specimen in excellent condition which 
was collected from the south-central Pacific, and, 
as noticed by Dr. Paxton, his specimen seemed to 
differ noticeably from Atlantic adults in having 
smaller scales, among other features. In 1975 Dr. 
Bruce Robison made available to us an adult col- 
lected in the Halmahera Sea during the R/V 
ALPHA HE Ix S.E.A.L. Expedition; he sub- 
sequently provided a second specimen from the 
same source. Recently Paxton informed us of an 
adult specimen in good condition collected by the 
R/V GALATHEA off Madagascar which he located 
in the Zoological Museum in Copenhagen. At last 
it was possible to compare an adult in good condi- 
tion from the western Indian Ocean (type locality 
of latifrons was off Zanzibar) with adults from 
the Pacific and with G. pumilus in the Atlantic 
and to evaluate the status of latifrons. 


ACKNOWLEDGMENTS 


We thank Dr. E. Bertelsen and officials of the 
Danish DANA Oceanographic Collections for 


making available four postlarval specimens and 
facilities for use by the senior author during a visit 
to Copenhagen in 1969. The National Marine 
Fisheries Service, Exploratory Fishing and Gear 
Research Base, Pascagoula, Mississippi, pro- 
vided most of the Atlantic adult specimens, par- 
ticularly through the efforts of Mr. Edward Klima 
and Mr. Bennie Rohr. Dr. John Paxton, Austra- 
lian Museum, Sydney, made available an adult 
specimen collected by the ORSTOM Center at 
Noumea, New Caledonia; Drs. Paxton and P. 
Fourmanoir (ORSTOM) kindly allowed us to uti- 
lize information in their unpublished manuscript 
on this specimen. Dr. Paxton also brought to our 
attention the existence of an adult specimen in 
the Zoological Museum of the University of 
Copenhagen collected by the GALATHEA off 
Madagascar; this specimen was loaned to us by 
Dr. Jorgen Nielsen. Dr. Bruce Robison, Univer- 
sity of California, Santa Barbara, made available 
two adults collected by the R/V ALPHA HELIx in 
the Halmahera Sea. Dr. Leslie Knapp, Smithso- 
nian Oceanographic Sorting Center, generously 
made available an adult from the western Indian 
Ocean. Mrs. Myvanwy Dick, Museum of Com- 
parative Zoology, Harvard University, loaned 
specimens. The staff of the U.S. National 
Museum of Natural History aided the junior au- 
thor while he was examining specimens in their 
care. Most postlarval specimens from the Atlan- 
tic resulted from the cooperation of the officers 
and crew of the University of Miami’s R/V 
PILLsBuRY. Dr. William M. Stephens greatly as- 
sisted in the early stages of preparation of this 
manuscript and with cinemaphotography of a liv- 
ing specimen. Mrs. Ann Jensen and Mr. Richard 
Schekter were especially helpful in assisting the 
senior author during PILLSBURY cruise P-6907. 
Mr. Schekter and Mr. Kenneth Kimball kindly 
examined stomach contents of Atlantic speci- 
mens. Mrs. Fay Mucha prepared histological sec- 
tions of an adult. Discussions about this study 
with Dr. C. Richard Robins were of much assis- 
tance. We are especially grateful to Ms. Joy God- 
frey for preparing the drawings of the fish. Drs. 
C. Richard Robins, Warren Frethofer, John Pax- 
ton, and Robert K. Johnson, and Mrs. Lillian 
Dempster offered suggestions on all or parts of 
early drafts of the manuscript. Mrs. Katherine 
Smith, Mrs. Terry Arambula Greenfield, Mr. 
James Gordon, Mr. Maurice Giles, Mr. Donald 
Heuer, Ms. Terry Loy, Mr. William Ruark, and 
Miss Pearl Sonoda aided in various ways. Mrs. 


DE SYLVA & ESCHMEYER: GIBBERICHTHYID FISHES 


Monica Abbott gave us valuable editorial assis- 
tance. 


METHODS 


The prejuvenile specimens are here termed the 
‘‘kasidoron”’ stage, although subsequent work- 
ers may wish to disregard this term. The kasido- 
ron stage is characterized by the presence of a 
unique pelvic appendage, and we confine the 
term to specimens possessing the appendage. 
The terminology ‘‘prejuvenile’’ comes from 
Hubbs (1958) as modified by Mansueti and Hardy 
(1967). 

The last two elements in the dorsal and anal fin 
are counted as one ray. The first elements in the 
dorsal and anal fin become spinous when the fish 
is between 20 and 30 mm standard length (SL). 
The fin-ray counts of adults are taken as the 
number of spinous points or spines plus soft rays 
(see text). 

Measurements are those commonly used to 
describe teleost fishes with a few modifications. 
Measurements originating from the anterior end 
of the fish are taken from the most anterior point 
of the left premaxillary. Head length is measured 
to the posteriormost tip of the upper opercular 
spine. Orbit diameter is the interior diameter of 
the bony orbit measured in a horizontal line. 
Measurements of prejuvenile specimens were 
made with an ocular micrometer. The third pelvic 
ray is measured to the tip of the ray without the 
pelvic tree, and the stalk of the pelvic tree is 
measured from the tip of the pelvic ray to the end 
of the main stalk. Many adults were variously 
damaged during capture, and some _ mea- 
surements are, therefore, inaccurate. Mea- 
surements of adults were usually taken to the 
nearest 0.5 mm. Original data used to prepare 
Figures 8-13 are on file at the California 
Academy of Sciences. 

Abbreviations of depositories of specimens are 
as follows: AMNH—American Museum of Nat- 
ural History, New York; AMS—Australian 
Museum, Sydney; ANSP—Academy of Natural 
Sciences of Philadelphia; BMNH—British 
Museum (Natural History), London; BOC— 
Bingham Oceanographic Collection, Yale Uni- 
versity; CAS—California Academy of Sciences, 
San Francisco; MCZ—Museum of Comparative 
Zoology, Harvard University; Sl1O—Scripps In- 
stitution of Oceanography, La Jolla; UMML— 
University of Miami, Rosenstiel School of 
Marine and Atmospheric Science, Miami; 


217 


USNM—United States National Museum of 
Natural History, Washington, D.C.; ZMUC— 
Zoological Museum, University of Copenhagen. 


MATERIAL EXAMINED 
(Number of specimens and their standard length [mm] in par- 
entheses.) 


Gibberichthys pumilus 

Kasidoron stage (depths of capture are as given by the col- 
lecting agency and are approximate estimates based on wire out 
and wire angle; these specimens were collected with 1-meter 
and 2-meter plankton nets): ANSP 102061 (1; 21.2, holotype of 
K. edom), 28°27' to 28°25'N, 73°42'W, 15 m over bottom of 4462 
m, PILLSBURY sta. 165, 2235-2335 hrs, 7 Aug. 1964. ANSP 
102062 (1; 15.7, paratype of K. edom), taken with the holotype. 
UMML 16213 (1; 7.8, paratype of K. edom), 32°46'N, 64°33'W, 
8 m over 2743 m, PILLSBURY sta. 144, 2025-2115 hrs, 3 Aug. 
1964. UMML 16214 (1; 12.1, paratype of K. edom), 28°30’ to 
28°27'N, 73°39’ to 73°42'W, 1-2 m over 4462 m, PILLSBURY Sta. 
164, 2115-2215 hrs, 7 Aug. 1964. UMML 28806 (1; 15.3), 
30°00'N, 68°00'W, 16 m over 5014 m, PILLsBURY sta. 1053, 
2109-2143 hrs, 24 Aug. 1969. ZMUC P41630 (1; 7.5), 21°50’N, 
50°12'W, 100 m wire out, DANA sta. 3543 IIT, 2145 hrs, 12 Aug. 
1928. ZMUC P41631 (1; 9.0), 24°48'N, 53°47’W, 17 m wire out, 
St. JAN sta. 543, 1930 hrs, 6 Aug. 1912. ZMUC P41632 (1; 11.1), 
24°10'N, 67°00'W, 77 m wire out, AG PETERSON sta. 803, 1900 
hrs, 21 Sept. 1914. 

Adults: CAS 14564 (2; 69 and 87, cleared and stained), 
T°41'N, 53°48’W, 677 m, OREGON II sta. 10606, 21-m shrimp 
trawl, 10 May 1969. CAS 14565 (3; 49, 53, and 71.5) and BMNH 
(1; 57.5), 29°16’N, 86°55’W, 640 m, OREGON II sta. 10913, 
58-m shrimp trawl, 12 Feb. 1970. FMNH 82924 (1; 80.5), 
7T°49'N, 54°22'W, 732 m, OREGON II sta. 10604, 21-m 
shrimp trawl, 10 May 1969. CAS 14566 (1; 91.5), 17°42'N, 
63°58’W, 741 m, OREGON II sta. 10832, 21-m shrimp trawl, 3 
Dec. 1969; CAS 37920 (1; 85), 7°41'N, 53°57’ W, 735 m, OREGON 
II sta. 10621, 30-m shrimp trawl, 16 May 1969. CAS 14567 (2; 63 
and 78) and CAS 14568 (1; 82.5, nerve preparation), 7°35'N, 
$3°29'W, 658 m, OREGON II sta. 10608, 39-m shrimp trawl, 11 
May 1969. CAS 14569 (1; 85.5), 29°09'N, 87°58’W, 841 m, 
OREGON II sta. 10650, 68-m shrimp trawl, 25 June 1969. CAS 
14570 (1; 83.5) and UMML 7041 (4; 57, 72.5 82.0, and 83.5), 
29°10'N, 87°55’ W, bottom depth 933-732 m (fishing depth un- 
certain), OREGON Sta. 2399, 12-m mid-water trawl, 4 Feb. 1959. 

The following additional adults were used in the map of 
distribution but were not examined in detail: MCZ 44211 (3 
specimens), 7°46'’N, 54°00’ W, 732 m, OREGON sta. 4299, 20-m 
shrimp trawl, 23 Mar. 1963. USNM 187664 (1), 29°54'N, 
80°11'W, 320-329 m, SILVER Bay sta. 3661, 24-m flat trawl, 16 
Jan. 1962. UMML uncat. (1; frozen), 20°40’ to 20°42'N, 
73°48’ to 73°40'W, est. depth 800-1100 m over 1463-2304 m, 
PILLSBURY sta. 1174, midwater trawl, 0441-0945 hrs, 29 June 
1970. USNM 187665 (1), 29°11'N, 87°47'W, 832 m, OREGON 
sta. 3218, 18/24-m semi-balloon trawl, 9 Feb. 1961. Plus litera- 
ture reference to one from OREGON sta. 1425, 29°04'N, 
88°05'W, 914 m, 24 Sept. 1955. Plus the following USNM uncat. 
lots: OREGON sta. 3654 (1), 29°08.5’N, 88°00.5’W, 732-750 m, 
12-m flat trawl, 25 July 1962. OREGON sta. 3660 (1), 29°10’N, 
87°57' W, 658-732 m, 12-m flat trawl, 27 July 1962. OREGON sta. 
3669 (1), 28°51.5'N, 88°39’ W, 622 m, 12-m flat trawl, 29 July 
1962. OREGON sta. 3670 (4), 29°00.5'N, 88°22'W, 732 m, 12-m 
flat trawl, 30 July 1962. OREGON sta. 4902 (2), 09°02.4'N, 
76°31.5'W, 732 m, 20-m shrimp trawl, 28 May 1964. Miss Vir- 
GINIA, no sta. (1), 29°10’N, 79°50’W, 180-200 m, Mar.—Apr. 


218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 6 


ee et 
AD fet \ 
f % i — P 
VIE MISS Ce é 
Sake va a AACAR ENN SSS 
ies \ \\ SSS OSS SS 
“ \ ; —— . — ~ _ = 
i ly ee ae en 
>" ‘ € ‘ ee 


‘ . * 
% 


Re A 


FiGure 2. Gibberichthys pumilus, kasidoron stage, 11.1 mm SL, ZMUC P41632. 


DE SYLVA & ESCHMEYER: GIBBERICHTHYID FISHES 


1952. Plus the specimens reported by Parr (1933): BOC 2838 (1; 
31.5, holotype of G. pumilus) 21°44'N, 72°43'25"W, 2134 m wire 
out, PAWNEE sta. 48, about 2127 hrs, 6 Apr. 1927. BOC uncat. 
(1, 91), 25°39'N, 77°18’ W, 1050 or 1100 m to surface, triangular 
midwater trawl, ATLANTIS sta. 1478, 20-21 Feb. 1933. 


Gibberichthys latifrons 

Kasidoron stage: ZMUC P41633 (1, 7.8), 2°17'N, 126°48.5’E, 
50 m of wire out, DANA sta. 3744 V, 2110 hrs, 7 July 1929. (Plus 
original description of K. latifrons, type locality off Zanzibar.) 

Adults: USNM 205551 (1; 112), 16°44’S, 43°44’E, 58 m, 
shrimp trawl, ANTON BRUUN cruise 8, sta. 407J, 15 Oct. 1964. 
ZMUC P41634 (1; 124), off Madagascar, 11°43’S, 49°09’E, 1300 
m, GALATHEA Expedition 1950-52, sta. 220, 1 Mar. 1951. AMS 
T.15999-001 (1; 103), 11°17’S, 142°47'W, 0-1040 m, 3-m IKMT, 
(ORSTOM) Corio_is sta. CARIDE III-17, 7-8 Feb. 1969. CAS 
37918 (1; 80), Halmahera Sea, 0°10.5'S, 128°33.3’E, 750-1000 
m, opening and closing 2.4-m Robison midwater trawl, ALPHA 
HELIx, S.E.A.L. Expedition, sta. 142, 1200-1400 hrs, 20 May 
1975. CAS 37919 (1; 89.6), Halmahera Sea, 0°08.9’S, 
128°40.0’E, 0-960 m, 2.4-m Robison midwater trawl, ALPHA 
HELIx, S.E.A.L. Expedition, sta. 137, 0955-1300 hrs, 19 May 
1975. 


COMPARISON OF KASIDORON EDOM WITH 
GIBBERICHTH YS PUMILUS 


Though a complete size series of specimens is 
not available, it is possible to show that Kasido- 
ron edom is the prejuvenile stage of Gib- 
berichthys pumilus. Evidence comes especially 
from metamorphosis of the pelvic appendage, 
examination of ontogenetic changes in the devel- 
opment of crests on the head, similarity of meris- 
tic features, and other characters. These are dis- 
cussed below. 


Pelvic appendage.—The most striking feature of 
the kasidoron stage is the presence of a peculiar 
pelvic appendage (pelvic arborescence of Robins 
and de Sylva 1965). This structure is described in 
detail by Robins and de Sylva (1965) and Robins 
(1966). The pelvic appendage is a highly modified 
third pelvic ray which, according to Robins and 
de Sylva (1965: 192), is **. . . fitted at its tip with a 
long, round stalk to which attach large, hollow, 
leaf-like sacs.’ The pelvic appendage they illus- 
trated was partially reconstructed and seems to 
have been an even larger arborescence (see also 
Thorp 1969, Fig. 1, for G. latifrons). Additional 
material available to us shows the development 
of this structure (Figs. 1-3). The pelvic tree is 
only partially developed in the 7.5-mm specimen 
(Fig. 1) and an 11.1-mm specimen (Fig. 2), and it 
then proliferates rapidly in size and complexity as 
shown in a 15.3-mm specimen (Fig. 3). (Further 
embellishment is found in the 21.0-mm holotype 


219 


of Kasidoron latifrons [Thorp 1969: Fig. 1].) The 
31.5-mm_ holotype of Gibberichthys pumilus 
lacks the pelvic appendage, as do all of our larger 
specimens. However, we do find evidence in 
adults which suggests the fate of the pelvic ap- 
pendage. In adults there is a discontinuity area 
part way up the third pelvic fin ray (Fig. 4) which 
is easily visible in cleared-and-stained speci- 
mens. We believe the expanded area represents 
the site of attachment of the pelvic appendage. It 
seems most likely that when the pelvic tree is lost 
the third pelvic ray continues to grow and seg- 
ment normally, producing the result shown in 
Fig. 4. This discontinuity occurs only on the third 
pelvic ray as evidenced by examination of three 
cleared-and-stained specimens. The third pelvic 
ray in most juveniles and adults shows some evi- 
dence of deformity; it is the thickest, and is fre- 
quently slightly bent or distorted. The pelvic ap- 
pendage is lost when the fish is between about 21 
and 31 mm standard length, the limits of our 
material which have or do not have the append- 
age. 


Body shape and coloration.—The 15.3-mm pre- 
juvenile specimen (Fig. 3), the 31.5-mm holotype 
of Gibberichthys pumilus (Fig. 5), and large 
adults (Figs. 6 and 7) show the changes in body 
shape with growth. Measurements (Figs. 8-13) 
reveal that there is a constant growth rate of most 
body parts when compared with standard length, 
but different slopes of the growth curves cause 
different apparent effects. For example, the orbit 
increases in diameter very slowly as the fish 
grows, so that the orbit is nearly one-half the head 
length in prejuvenile specimens but one-seventh 
or one-eighth of the head length in adults. Mea- 
surements of some body parts when plotted 
against standard length show slight changes in 
growth rates at about a size where the pelvic 
appendage is lost, and presumably when the 
juvenile specimens move to a deeper habitat. For 
example, the snout length (Fig. 11) grows slowly 
in prejuvenile specimens, subsequently increases 
rapidly, and then shows a slower constant growth 
rate in adults. Head length (Fig. 13) seems to 
show a similar change in growth rate. The length 
of the third pelvic fin ray (Fig. 12) increases very 
slowly, thereby becoming very small propor- 
tionally in larger specimens, but some variability 
in our measurements results from the third ray 
being broken in large specimens; the develop- 
ment of the pelvic appendage (Fig. 12) is ex- 


220 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 


FiGuRE 3. Gibberichthys pumilus, kasidoron stage, 15.3 mm SL, UMML 28806. 


tremely rapid. Other measurements are given in 
Figs. 8-13. 

The most notable changes in body shape, be- 
sides proportional differences in eye size, are the 
shift in position of the mouth from an oblique 
position to a more terminal one, and a change in 


the orbit from a sub-superior to a more lateral 
position. The proliferation of thin, spiniferous 
crests over the head of the prejuveniles and 
juveniles is accompanied by closure of the 
cavernous sulci permeating the head. This transi- 
tion is observed in the fenestration of the su- 


FiGure 4. Lateral view of the third pelvic fin ray of adult Gibberichthys pumilus. Arrow indicates discontinuity (see text). 


nN 
nN 
— 


DE SYLVA & ESCHMEYER: GIBBERICHTHYID FISHES 


FicurE 5. Holotype of Gibberichthys pumilus, 31.5 mm SL, after Parr, 1933. 


— 


(he 


FiGuRE 7. Gibberichthys pumilus, 87 mm SL, CAS 14564, cleared and stained. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 6 


5 ha ha) 


Orbit diameter 
Length of pectoral fin e 
Jaw length o 


PART IN mm 


0 10 20 30 40 50 60 70 80 90 100 110 120 


STANDARD LENGTH mm 
Ficure 8. Orbit diameter, length of pectoral fin, and jaw length in Gibberichthys pumilus and G. latifrons (enlarged symbols) 


Snout to origin Ist dorsal spine * 
Snout to origin Ist dorsal ray o 


Snout to Ist anal spine @ 


PART IN mm 


80 90 100 10 120 


0 10 20 30 40 50 60 70 
STANDARD LENGTH mm 


Ficure 9. Lengths of snout to origin of first dorsal spine, snout to origin of first dorsal soft ray, and snout to origin of first anal 
spine in Gibberichthys pumilus and G. latifrons (enlarged symbols). 


DE SYLVA & ESCHMEYER: GIBBERICHTHYID FISHES 


20 


Interorbital width * 
Longest dorsal ray @ 
Longest anal ray o 


PART IN mm 


0 0 20 30 40 50 


60 70 80 90 100 10 120 


STANDARD LENGTH mm 


Ficure 10. Interorbital width, longest dorsal soft ray, and longest anal soft ray in Gibberichthys pumilus and G. latifrons 


(enlarged symbols). 


praorbital region of a 15.3-mm specimen (Fig. 14a) 
and is also visible in the same area in adults (Fig. 
14b). Similarly, the bony crests so characteristic 
of the head of adults (Fig. 6-7, 14) are just ob- 
servable in the internasal region of a 15.3-mm 
specimen (Fig. 15). As growth continues, the rel- 
ative position of several cranial bones changes 
concomitantly to correspond with the lengthen- 
ing and flattening of the head. These include the 
change in position of the maxillary to accommo- 
date a rearward shift in the angle of the preoper- 
cle and opercle. The maxillary is unsheathed in 
prejuveniles, but growth of the circumorbital 
bones partially sheathes the maxillary in larger 
specimens. 

The height of the lumps on the midline behind 
the head (one character used by Thorp (1969) to 
distinguish K. latifrons from K. edom) is due to 
three interneural (predorsal) bones (see espe- 
cially Figs. 3, 6-7; see Thorp, 1969, Fig. 1, for G. 
latifrons). Their manifestation as lumps seems 
more prominent in the prejuveniles than in 


adults, but this variability may also result from 
shrinking with preservation. 

Little ontogenetic change in coloration occurs; 
both juveniles and adults are black. The eyes of a 
freshly captured adult (PILLSBURY sta. 1174) 
glowed red when a flashlight was shined on them. 


Meristic characters.—Counts of dorsal, anal, 
pelvic, and pectoral rays are similar both in the 
kasidoron stage and in adults (Table 1). The an- 
terior fin elements in the dorsal and anal fins 
become spinous when the fish reaches a size be- 
tween our largest kasidoron stage (21.2 mm) and 
the holotype of Gibberichthys pumilus (31.5 
mm). The anterior spines are fixed on broad, firm 
bases; usually only the last spine is movable. 
There are usually 6 spinous points in the dorsal 
fin and 4 in the anal fin. The first plate of the anal 
fin has 2, while subsequent plates each bears | 
spinous point. Our adult specimens average 
about 1% ‘‘ray’’ higher in dorsal ‘*fin-ray’’ count 
when spinous points plus soft rays of adults are 


224 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 6 


Snout length 
Depth of caudal peduncle o 
Length of caudal fin @ 


PART IN mm 


60 70 80 90 100 10 120 


STANDARD LENGTH mm 
FiGureE 11. Snout length, depth of caudal peduncle, and length of caudal fin in Gibberichthys pumilus and G. latifrons 


(enlarged symbols). 


compared with fin rays of prejuveniles. We be- 
lieve the first plate of the dorsal and anal fin 
probably forms from fusion of two fin rays, but 
usually an extra spinous point develops on the 
first plate of the dorsal fin which results in the 
higher dorsal ‘‘fin-ray’’ count of adults. (If three 
fin rays make up the first plate when it bears three 
spinous points, then at least one of these ele- 
ments is not distinguishable in prejuveniles.) 
Pelvic rays are usually 6, with the first ray 
spinous in adults. One specimen, the 7.8-mm 
paratype of K. edom, had only 5 countable pelvic 
elements, as did one adult. Procurrent caudal 
rays become spinous after the kasidoron stage; 


Third pelvic ray * 
Stalk @ 


PART IN mm 


they usually number 7 above and 6 below. Princi- 
pal caudal rays usually number 19, with one un- 
branched above and one below. The caudal fin 
structure of an adult is illustrated in Fig. 16. 

Vertebrae number 29-30 in three postlarvae 
and 20 adults of G. pumilus examined for this 
character. 


Pyloric caeca.—Pyloric caeca number 12-13 in 
adult specimens examined for this feature with 
about 9 exposed and 3 or 4 under the liver; most 
caeca form one whorl, but some caeca lie farther 
down the intestine. In the holotype of K. edom 
there were 9 caeca arranged in a single whorl 


STANDARD LENGTH mm 


Ficure 12. Length of third pelvic fin ray and length of the pelvic appendage in Gibberichthys pumilus and G. latifrons 


(enlarged symbols). 


DE SYLVA & ESCHMEYER: GIBBERICHTHYID FISHES 


Head length » 
Greatest body depth o 
Depth at Ist dorsal spine @ 


Depth at Ist anal spine x 


PART IN mm 


40 50 


60 


70 80 120 


STANDARD LENGTH mm 


FiGure 13. Head length, greatest body depth, body depth at origin of first dorsal spine, and body depth at origin of first anal 
spine in Gibberichthys pumilus and G. latifrons (enlarged symbols). 


(Robins and de Sylva, 1965: 196), but possibly 
some caeca were rubbed off during examination 
of this very small specimen, or they were not yet 
developed. In the prejuvenile specimens and in 
adults the caeca are pale and long. 


Gill rakers.—Counts of gill rakers are particu- 
larly difficult to make on the prejuvenile speci- 
mens, but the number of rakers on the first arch in 
these specimens and in adults (Table 1) are com- 
parable. Counts for adults are 5—6 above, one at 
the angle, and 13-15 on the lower arch, total 
18=22)\ (mean — 20:2) Ni = 15). 


Lateral line.—The lateral line in type specimens 
of K. edom was reported as composed of about 32 
vertical rows of raised papillae (Robins and de 
Sylva 1965: 195). Most skin on the body was 
rubbed off during caputre of our adult specimens, 
but in some specimens sufficient skin is intact to 


permit us to determine that adults also have a 
lateral line composed of vertical rows of raised 
pores, each row with about 6 to 8 pores. We are 
unable to give an exact count of the number of 
vertical rows, but they are spaced approximately 
as in the holotype of K. edom. Parr (1934: Fig. 37) 
illustrated the vertical rows of pores and found 28 
rows in his 91-mm specimen. 


Swimbladder.—Robins and de Sylva (1965: 196) 
reported that K. edom has a well developed 
swimbladder restricted to the anterior portion of 
the body cavity. The swimbladder in adult 
specimens is moderate, thin walled, and lies in 
the anterior half or two-thirds of the body cavity. 
It is partially filled with fat in adults. 

In summary, differences in the above features 
can be attributed to ontogenetic changes, fea- 
tures which are the same or nearly so in the small 
specimens and adults, and those differences 


226 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 6 


"5 Taian 


FiGcure 14. Lateral view of oribital region in (A) postlarva, 
UMML 28806, 15.3 mm SL, and (B) small adult, UMML 7401, 
72.5 mm SL, of Gibberichthys pumilus. 


which may result from error because of the small 
size of prejuveniles examined. We believe that 
there can be no doubt that Kasidoron edom is the 
prejuvenile stage of Gibberichthys pumilus. 


SPECIES OF THE GENUS 
GIBBERICHTHYS 


Since Kasidoron edom is the prejuvenile of 
Gibberichthys pumilus , then Kasidoron latifrons 
Thorp from the western Indian Ocean is a pre- 
juvenile of the same species or a separate species. 
We have available only six specimens from out- 
side the Atlantic; Ebeling and Weed (1973) re- 


rr 


Tuo = 


FiGure 15. Dorsal view of nasal region in (A) postlarva and 
(B) a small adult of Gibberichthys pumilus (same specimens 
as in Fig. 14). 


ported one non-Atlantic specimen, and the 
holotype of /atifrons brings to eight the total 
non-Atlantic specimens known. 

The differences between the holotype of latif- 
rons and prejuveniles of pumilus as given by 
Thorp (1969) are as follows: the lumps behind the 
head are larger and the body deeper in Jatifrons , 
and /atifrons has a minute extra pore in the head 
lateralis system, no small cycloid scales along the 
lateral line, a lower gill-raker count, slight differ- 
ences in measurements, and differences in the 
pelvic appendage. The dorsal lumps are caused 
by three interneural (predorsal) bones, and the 
prominence of the lumps is probably related to 
some extent by condition or shrinkage with pres- 
ervation. We are unable to assess the minute 
extra head pore. The number of gill rakers and 
other counts (Table 1) fall within the range of 
pumilus based on our material. We are unable to 
provide much additional information on differ- 
ences in the pelvic appendage (see Thorp 1969: 
68-69) but suspect this to be a structure which is 
variable in details; no Atlantic prejuvenile had 


DE SYLVA & ESCHMEYER: GIBBERICHTHYID FISHES 


227 


TABLE |. COUNTS FOR SPECIMENS OF Gibberichthys pumilus AND G. latifrons. 


Dorsal fin rays 


Prejuveniles Spinous points + rays in adults Total (Adults) 
Liss 4 15 5 6 W 8 + 8 9 IS ees 1G Oy: 
G. pumilus' 4 3 I l 6 9 - - 5, 10 De) WE GS ee 
G. latifrons® 1 = | = - 1 3 1 = 5 = = l 3 l 
Anal fin rays 
Prejuveniles Spinous points + rays in adults Total (Adults) 

10 IL] 12 4 5) + q 8 y) 11 2 13 14 

G. pumilus' 2 2 5 13 oy} a if l 6 7 2 — 
Gilatijrons= = 2 = = 5 2 2, l = 2 2 l 

Pectoral rays Pelvic rays® 
(left side) (left side) Gill rakers? (left side) 

13 14 15 > 6 18 19 20 21 up) 

G. pumilus' 4 16 4 2 74) 3 3 6 4 5 
G. latifrons? l 6 = - 7 l 3 — - 2 


' Includes Parr’s (1933) count for one specimen. 
* Includes Thorp’s (1969) count for holotype. 
3 Some small specimens not included. 


the arborescence as well developed as Thorp’s 
specimen, and the differences in pigmentation of 
the “‘leaves’’ as presented by Thorp may prove to 
be real differences. 

Comparison of the five non-Atlantic adults 
with the larger series of Atlantic adults reveals 
virtually no differences between them in counts 
and measurements (see Table 1; Figs. 8-13). Our 
largest specimen of G. pumilus is 91.5 mm in 
standard length while the largest specimen of G. 
latifrons is 124mm SL. One major difference was 
pointed out by Dr. John Paxton when he for- 
warded the AMS specimen to us. The scales on 
the body appeared to be considerably smaller in 
his specimen from the southwestern Pacific than 
in the specimen figured by Parr (1933). The skin 
and scales are more intact in our non-Atlantic 
specimens and all of the Atlantic adults are miss- 
ing virtually all scales and much skin. We esti- 
mate that the scales in Atlantic specimens are 
about twice the size of scales in the non-Atlantic 
specimens. This difference is easily seen by com- 
paring Parr’s figure (1933: Fig. 1; reproduced 
here as Fig. 5) and our Figure 6, with the excellent 
rendition of an adult /atifrons in Ebeling and 
Weed (1973: Fig. 12). (Parr’s illustration showing 
a row of enlarged scales along the lateral line 


probably is inaccurate, and we believe that two 
slightly enlarged scales are present, rather than 
one enlarged and elongated (vertically) scale, and 
that both scales house papillae of the lateral line.) 

Besides the difference in scale size, the Atlan- 
tic and non-Atlantic specimens differ in vertebral 
counts as given below: 


Vertebrae Pio es) SV oss 
Atlantic specimens Sse = 
Non-Atlantic specimens - —- 2 3 


Because of certain differences in prejuvenile 
features, and the differences in scale size and 
number of vertebrae, we recognize two species, 
G. pumilus from the Atlantic and G. latifrons 
from the western Pacific and Indian oceans. 


SYNONYMIES 


Prior literature may be summarized as follows: 
Family GIBBERICHTHYIDAE Parr 


Gibberichthyidae Parr, 1933: 1, 4-5 (type-genus Gibberichthys 
Parr, 1933; comparison with other beryciform families). 
Ebeling 1962: 11 (in Stephanoberyciformes). Ebeling and 
Weed: 1973. 397 et seq. (relationships). Greenwood et al. 
1966: 398 (listed in order Beryciformes, suborder 
Stephanoberycoidei). Rosen and Patterson 1969: 456, 461 


228 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 6 


FiGurRE 16. Caudal skeleton of Gibberichthys pumilus ,CAS 
14564, 87 mm SL. 


(suggested Kasidoron was young of Gibberichthys; relation- 
ships). Rosen 1973: 469 et seq. (relationships). 
Kasidoroidae Robins and de Sylva, 1965: 190 et seq. (type- 
genus Kasidoron Robins and de Sylva, 1965). Robins 1966: 
696-701 (descriptive information, relationships). 
Kasidoridae (sic), Myers and Freihofer 1966: 193-194 (com- 
pared with Megalomycteridae). Greenwood et al. 1966: 395 
(listed in suborder Mirapinnatoidei, order Cetomimiformes). 


Genus Gibberichthys Parr 


Gibberichthys Parr, 1933: 5 (type-species Gibberichthys 
pumilus Parr, by original designation, monotypic). 

Kasidoron Robins and de Sylva, 1965: 190 (type-species 
Kasidoron edom, by original designation, monotypic). 


Gibberichthys pumilus Parr 


Gibberichthys pumilus Parr, 1933: 5—6, Fig. 1 (type-locality 
21°44'N, 72°43'25”W, 2134 m wire, 6 Apr. 1927, PAWNEE Sta. 
48: BOC 2838). Parr 1934: 35-36, Fig. 11 (description of 
9l-mm specimen; ATLANTIS sta. 1478; compared with 
holotype). Myers 1936: 118 (belongs near Stephanoberyci- 
dae). Springer and Bullis 1956: 66 (listed; Gulf of Mexico, 
OREGON sta. 1425). Grey 1959: 333-334 (description of 
a 47-mm specimen, 29°04’N, 88°05’'W, 914 m, OREGON 
sta. 1425). Ebeling and Weed 1973: 412-414 (in part; 
figure and part of description is G. latifrons; family relation- 
ships). 

Gibberichtys pumilus, Ebeling 1962: 11 (misspelled genus; 
agreed with Rofen that it was allied to melamphaeids). 

Kasidoron edom Robins and de Sylva, 1965: 189 et seq., Figs. 
1-2 (original description; types from near Bermuda and NE 
of Bahamas). Robins 1966: 696 et seq., Figs. 1-3 (additional 
information on K. edom; relationships). Voss and Sisson 


1967: 393 (figure). Thorp 1969: 62 et seq. (compared with K. 
latifrons). Rass 1971: plate opposite p. 97 (figure; tips of 
pelvic appendage shown as luminescent). 


Gibberichthys latifrons Thorp 


Gibberichthys latifrons Thorp, 1969: 61-70, Figs. 1 (original 
description; type-locality near Zanzibar, 08°34'S, 41°37’E; 
good description; compared with K. edom). 

Gibberichthys pumilus , Ebeling and Weed 1973: 412-414, Fig. 
12 (good figure; did not distinguish /atifrons from edom). 


DISTRIBUTION AND BIOLOGY 


Gibberichthys pumilus is now known from 
several localities in the western Atlantic (Fig. 17). 
Gibberichthys latifrons is known from the west- 
ern Indian Ocean (3 specimens), the Halmahera 
Sea area (3), near Samoa (Ebeling and Weed 
1973) (1), and west of the Marquesas Islands (1). 
We attribute the separate distribution of pre- 
juveniles and adults in the Atlantic Ocean (Fig. 
17) to an artifact of collecting methods and effort. 
The prejuveniles were captured in oceanic waters 
where University of Miami PILLSBURY opera- 
tions were concentrated, and most adults were 
caught in coastal waters where OREGON and 
OREGON II trawling operations were conducted. 
We expect the species is widespread in the west- 
ern Atlantic. 

Lack of data from closing nets allows us only 
limited interpretation of the vertical distribution 
of adults. The prejuveniles are epipelagic, all cap- 
tures being taken between near-surface waters 
and about 50 m. All captures were made at night 
and no specimens were taken in near-surface day- 
light tows. One might infer from the black color- 
ation of the fish that prejuveniles may undergo 
diel migration at least into lower epipelagic or 
upper mesopelagic levels during daylight hours. 
Changes in morphology between the prejuvenile 
stage and the juvenile stage, which occur at about 
the time that the fish descend into the lower 
mesopelagic or upper bathypelagic levels, are 
discussed in an earlier section. 

Capture of adults has been by use of both bot- 
tom trawls and midwater nets. The shallowest 
bottom trawl haul in the western Atlantic was in 
320 m and the deepest in 841 m, with most in the 
range of 650-750 m. Additional collections of 
adults from vertical tows and non-closing midwa- 
ter nets add limited information; one capture was 
estimated to be at 800-1100 m and another in 1050 
or 1100 m. It would appear that the depth range of 
adults in the western Atlantic is roughly from 300 


DE SYLVA & ESCHMEYER: GIBBERICHTHYID FISHES 


= Kasidoron 
Adults 


stage 


229 


FiGure 17. Distribution of Gibberichthys pumilus in the western Atlantic Ocean (symbols represent stations). 


m to 1000 m; however, one adult of /atifrons from 
the western Indian Ocean was collected at a 
depth of 58 m (correspondence with Leslie 
Knapp indicates that the depth of capture is accu- 
rate). The adult from the southwestern Pacific 
was from an oblique tow between 0 and 1040 m, 
and the one reported by Ebeling and Weed (1973) 
between 0 and 1000 m. The presence of fat in 
swimbladders of adults and the absence of cap- 
tures in near-surface waters at night, where 
most trawl hauls for midwater fishes have been 
made, suggests that adults do not undergo ver- 
tical migration, at least not into near-surface 
waters. 

The food of postlarval specimens was reported 
by Robins and de Sylva (1965: 200) as copepods, 
particularly calanoid copepods. Stomachs of 16 
adults of Gibberichthys pumilus were removed 
and the contents examined. Fifteen specimens 
contained food, 13 of which had identifiable con- 
tents as follows: 


Number of 
stomachs in which 


Category organism was found 


Amphipoda 
Gammaridea 
Lysianassidae 
Cyphocaris johnsoni 1] 
Hyperiidea 
Hyperiopsidae 
Hyperiopsis sp. 1 
Copepoda 
Calanoida 
Eucalanidae 
Eucalanus elongatus 1 
Metridiidae 
Pleuromamma abdominalis 1 
Harpacticoida 1 


The pelagic gammaridean Cyphocaris johnsoni 
was by far the most common food of adults, oc- 
curring in 11 stomachs, usually from | to 8 per 
stomach. Nearly all stomachs contained crusta- 
cean fragments, usually of amphipods. Accord- 
ing to Vinogradov (1970: 209) the pelagic gam- 


230 


marids in the total plankton mass above 3000 m is 
negligible, but since these amphipods were com- 
mon in stomachs and because other plankters 
were few, we suspect that Gibberichthys pumilus 
is selective in its food habits. 

The use of the pelvic appendage deserves addi- 
tional study. Robins and de Sylva (1965: 199) 
reported on the swimming behavior of a freshly 
captured specimen in a shipboard aquarium. 
They suggested that the pelvic appendage may 
mimic the nectosome of siphonophores. A more 
complete description of the pelvic appendage 
was given by Robins (1966: 698-700), particularly 
regarding microstructure. No sign of any spe- 
cialized structure that could account for lumines- 
cence was found, and no muscle fibers or nerves 
were found associated with the stalk or ‘‘leaves.”’ 
However, the epithelium on the tips of the pelvic 
appendage had been sloughed off in the specimen 
examined by Robins and de Sylva (1965) (per- 
sonal communication from Mrs. Priscilla Ras- 
quin Breder), so that epithelial luminescent struc- 
tures could have been lost. A second specimen 
was observed alive by de Sylva (PILLSBURY Sta. 
P-1053). The specimen lived for about four hours, 
swam only moderately, alternating bursts of ac- 
tive swimming near the surface with periods of 
quiet on the bottom. Its behavior appeared ab- 
normal. Periodic examination of the specimen in 
the dark showed that the ‘‘leaf’’ tips, if lumines- 
cent, were not noticeably so, but the room was 
not completely dark. The tips of the fresh 
‘leaves’ were pale greenish yellow, a cast some- 
times characteristic of luminous structures (see 
Nicol 1967, 1969). A color illustration which 
shows the presumed luminous nature of the 
epithelium of the tips is presented in Rass (1971: 
plate opposite p. 97). The simple nature of the 
pelvic appendage would seem to rule out volun- 
tary control of the luminescence, but bacterial as- 
sociations might be possible. (However, we 
would caution against illustrations depicting 
luminescence [e.g., Rass 1971] because it is quite 
possible that no luminescence is involved.) The 
apparent lack of muscles associated with the pel- 
vic appendage renders it passive and would seem 
to limit its use to either mimicry as proposed by 
Robins and de Sylva (1965) or general deception, 
particularly when viewed from below or behind, 
or as a “‘cover’’ source which might attract food 
items. The arborescence in Thorp’s specimen 
(1969: Fig. 1) resembles Sargassum weed more 
than in Robins’ specimen (1966: Fig. 1). 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 6 


Ripe gonads were found in two female speci- 
mens measuring 81 and 87 mm SL collected on 
May 10th. The eggs contain an oil globule. There 
is no evidence that the species is hermaphroditic. 


REMARKS 

The relationships of the family Gib- 
berichthyidae deserve additional study. We 
agree with Ebeling and Weed (1973) that the fam- 
ily Gibberichthyidae seems to show closest rela- 
tionships to the family Stephanoberycidae, and a 
more distant relationship to the Melamphaeidae. 
A brief examination of stephanoberycids and 
melamphaeids was made, and we found no indi- 
cation that a prejuvenile pelvic appendage exists 
in any xenoberyciform group except Gib- 
berichthys. Some melamphaeids, particularly 
postlarval specimens of the genus Poromitra, 
have extremely long pelvic fins with the indi- 
vidual rays branched many times and becoming 
hair-like distally. The family Gibberichthyidae is 
uniquely characterized by the presence of the 
pelvic appendage in postlarval specimens. It was 
separated by Norman (1957: 213) from melam- 
phaeids and stephanoberycids on the basis of the 
presence of spines in the pelvic, dorsal, and anal 
fins in Gibberichthys , especially the rigid spines 
with broad, flattened basal plates in the dorsal 
and anal fins (see also Ebeling and Weed 1973). 
When one recalls that the soft fin rays of the 
kasidoron stage become the fin spines of Gib- 
berichthys , it is not difficult to consider that simi- 
lar transitions of rays to fin spines may also occur 
in other groups of deep-sea fishes which are sus- 
pected of bizarre metamorphoses. 

We also briefly examined specimens of Ron- 
deletia bicolor and were struck by the re- 
semblance of this fish to Gibberichthys, as was 
Rofen (see Ebeling 1962: 11; Ebeling and Weed 
1973: 399). Ebeling and Weed (1973: 399) dis- 
cussed the problems surrounding placement 
of Rondeletia and the classification of three 
groups, their orders Xenoberyces (Stephano- 
beryciformes), Beryciformes, and Cetunculi 
(Cetomimiformes). Add to this the fact that 
Robins (1966) placed the postlarval Kasidoron 
edom in the Mirapinniformes. In turn he placed 
these near the Cetomimiformes, and noted 
that some of them were prejuveniles of ceto- 
mimids (Robins 1974), e.g. Megalomycter, Atax- 
olepis (C. R. Robins, personal communica- 
tion). However, Rosen and Patterson (1969) re- 
moved Kasidoron to the Beryciformes (including 


DE SYLVA & ESCHMEYER: GIBBERICHTHYID FISHES 


also the Cetomimoidei) but did not comment on 
the remainder of the mirapinniform fishes except 
to include them (op. cit.: 461) in the Lam- 
pridiformes. Nor does the reshuffling and reor- 
ganizing of these groups by Rosen and Patterson 
(1969) from the classification of Greenwood et al. 
(1966) advance our understanding of relation- 
ships except to move the Lampridiformes next to 
the Beryciformes and place all of them in the 
same superorder. Rosen (1973) has further re- 
shuffled these and other groups and presented 
(op. cit.: 469) a working hypothesis that there is a 
group (iv) within a broad order Beryciformes 
containing the Barbourisiidae, Rondeletiidae, 
Gibberichthyidae, Cetomimidae, Mirapinnidae, 
Eutaeniophoridae, Megalomycteridae, Melam- 
phaeidae, Stephanoberycidae, and Anoplogas- 
teridae. It is apparent that the higher-category 
classification of these fishes remains uncertain. 

Recent collections of deep-water fishes by 
many researchers are disclosing that our knowl- 
edge of metamorphosis of deep-water fishes is 
poorly known, and bizarre transformations in- 
volving more than one metamorphosis may be 
common in certain fish groups. Certainly, the 
identity of such groups as the Rosauridae, 
Megalomycteridae, Mirapinnidae, and Eutae- 
niophoridae should be re-examined, and some 
are being studied by other workers. It is possible 
that at least some of these represent prejuvenile 
stages of cetomimoid, berycoid, or other fishes. 


LITERATURE CITED 


EBELING, ALFRED W. 1962. Melamphaidae I. Systematics 
and zoogeography of the species in the bathypelagic fish 
genus Melamphaes Gunther. Dana Rep. Carlsberg Found. 
58: 1-164, figs. 1-73. 

AND WALTER H. WEED, III. 1973. Order 
Xenoberyces (Stephanoberyciformes). Jn Fishes of the 
western North Atlantic. Mem. Sears Found. Mar. Res., Yale 
Univ. | (pt. 6): xix + 698 pp. 

GREENWOOD, HumpHrRY P., DONN E. ROSEN, STANLEY H. 
WEITZMAN, AND GEORGES. MYERS. 1966. Phyletic studies 
of teleostean fishes, with a provisional classification of living 
forms. Bull. Am. Mus. Nat. Hist. 131(4): 339-456, pls. 
21-23. 

GREY, MARION. 1959. Deep sea fishes from the Gulf of 
Mexico with the description of a new species Squalogadus 
intermedius (Macrouridae). Fieldiana, Zool. 39(29): 323-346, 
figs. 54-57. 

Husss, Cart L. 1958. Dikellorhynchus and Kanazawaich- 
thys: nominal fish genera interpreted as based on pre- 
juveniles of Malacanthus and Antennarius, respectively. 
Copeia 1958(4): 282-285. 

MANSUETI, ALICE J., AND JERRY D. HARDY, JR. 1967. 
Development of fishes of the Chesapeake Bay region. An 


Nm 
Ww 


atlas of egg, larval, and juvenile stages. Part 1. Natural Re- 
sources Inst., Univ. Maryland. 202 pp. 

Myers, GeorGe S. 1936. A note on the stephanoberycid 
fishes. Copeia 1936(2): 118. 

, AND WARREN C. FREIHOFER. 1966. Megalomyc- 
teridae, a previously unrecognized family of deep-sea 
cetomimiform fishes based on two new genera from the 
North Atlantic. Stanford Ichthyol. Bull. 8(3): 194-206, figs. 
1-5. 

NicoL, J. A. C. 1967. The luminescence of fishes. Symp. 
Zool. Soc. Lond. 19: 27-55, 14 figs. 

1969. Bioluminescence. Pp. 355-394 in Hoar, S. and 
D. Randall, eds., Fish physiology, vol. 3. Reproduction and 
growth: bioluminescence, pigments, and poisons. Academic 
Press, New York, 485 pp. 

NorMan, J. R. 1957. A draft synopsis of the orders, families, 
and genera of Recent fishes and fish-like vertebrates. The 


British Museum (Natural History), London. 649 pp. 
[Copyrighted 1966.] 
ParR, ALBERT E. 1933. Deepsea Berycomorphi and Per- 


comorphi from the waters around the Bahama and Bermuda 

islands. Bull. Bingham Oceanogr. Collect. Yale Univ. 3(6): 

I-51, figs. 1-22. 

. 1934. Report on experimental use of a triangular trawl 
for bathypelagic collecting, with an account of the fishes 
obtained and a revision of the family Cetomimidae. Bull. 
Bingham Oceanogr. Collect. Yale Univ. 4(6): 1-59, figs. 
1-21. 

PATTERSON, COLIN. 1969. The caudal skeleton in Mesozoic 
acanthopterygian fishes. Bull. Br. Mus. (Nat. Hist.), Geol. 
17(2): 1-102, 28 figs. 

Rass, T. S. (editor). 1971. The life of animals. Vol. 4, no. 1, 
Fishes [in Russian]. Izdatel’stvo Prosveshcheniye, Mos- 
cow. 655 pp., 54 pls. 

Ropins, C. RICHARD. 1966. Additional comments on the 
structure and relationships of the mirapinniform fish family 
Kasidoroidae. Bull. Mar. Sci. 16(4): 696-701, figs. 1-3. 

. 1974. Review of ‘Fishes of the western North Atlan- 

tic. No. 1, part six. Order Heteromi (Notacanthiformes), 

Suborder Cyprinodontoidei, Order Berycomorphi (Be- 

ryciformes), Order Xenoberyces (Stephanoberyciformes), 

Order Anacanthini (Gadiformes) in part Macrouridae.”’ 

Copeia 1974(2): 574-576. 

, AND DONALD P. DE SYLVA. 1965. The Kasidoroidae, 
a new family of mirapinniform fishes from the western North 
Atlantic Ocean. Bull. Mar. Sci. 15(1): 189-201, figs. 1-2. 

RosEN, DONN Eric. 1973. Interrelationships of higher 
euteleostean fishes. Zool. J. Linn. Soc., Suppl. 1, 53: 397- 
513. 


, AND COLIN PATTERSON. 1969. The structure and 
relationships of the paracanthopterygian fishes. Bull. Am. 
Mus. Nat. Hist. 141(3): 359-474, pls. 52-78, 78 figs. 

SPRINGER, STEWART, AND HARVEY R. BULLIS, JR. 1956. Col- 
lections by the OREGON in the Gulf of Mexico. U.S. Fish 
Wildl. Serv., Spec. Sci. Rep. Fish. 196. 134 pp. 

THorp, C. H. 1969. A new species of mirapinniform fish 
(family Kasidoroidae) from the western Indian Ocean. J. 
Nat. Hist. 3(1): 61-70, 4 figs. 

VINOGRADOV,M.E. 1970. Vertical distribution of the oceanic 
zooplankton. Tr. Inst. Okeanol., Akad. Nauk SSSR. 
[Translated from Russian by the Israel Program for Scientific 
Translation, Keter Press, Jerusalem. 339 pp.] 

Voss, GILBERT L., AND ROBERT F. Sisson. 1967. Squids: 
jet-powered torpedoes of the deep. Natl. Geogr. Mag. 131(3): 
386-411, 29 figs. 


1 ie I 
C253X 
NH 


PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 7, pp. 233-251; 11 figs.; 5 tables 


August 18, 1977 


A NEW GENUS (MESOBIUS), AND THREE NEW BATHYPELAGIC 
SPECIES OF MACROURIDAE (PISCES, GADIFORMES) FROM 
THE PACIFIC OCEAN 


By 


Carl L. Hubbs 
Scripps Institution of Oceanography, University of California, San Diego, La Jolla, California 92093 


and 


Tomio Iwamoto 


California Academy of Sciences, San Francisco, California 94118 


ABSTRACT: A new genus, Mesobius, and three new species (M. berryi, M. antipodum, Nezumia parini) of 
bathypelagic macrourid fishes are described from the Pacific Ocean. M. berryi is known from the eastern 
North Pacific, M. antipodum from a single adult off New Zealand, and N. parini from the equatorial eastern 
Pacific and midwaters between Panama and northern Chile. M. berryi passes through a phalacromacrurus 
stage characterized in early prejuveniles by bold spotting on the head and body, stylopterus pectorals, and 
almost vertical jaws. Later prejuveniles have normal pectorals and more nearly horizontal mouths. Spots 
remain on the trunk and tail in individuals as large as 40 mm in head length, but the general blackish color 
characteristic of all bathypelagic macrourids obtains in older individuals. The small jaw teeth in narrow 
bands, the uniquely elongated and aligned head scales, and the seven branchiostegal rays of Mesobius 
contrast with the fanglike teeth in single series, the circular head scales, and the six branchiostegal rays of two 
other bathypelagic macrourids, Odontomacrurus murrayi and Cynomacrurus piriei. It appears to be most 
closely related to the monotypic genus Echinomacrurus. Nezumia parini, the only known bathypelagic 
member of an otherwise exclusively benthopelagic genus, retains many of the characters of its benthopelagic 
relatives, but it has the reduced musculature, skeleton, gill-filaments and gas-bladder of other bathypelagic 
macrourids. Its small scales are distinctively covered with very fine, erect spinules, that yield a velvety texture 
to body surface. 


INTRODUCTION 


Most of the 250 or so species of the deepsea 
fish family Macrouridae are benthopelagic as 
adults. These benthopelagic species generally 
hover and glide over the bottom at an inclined, 
nose-downward attitude while seeking out their 
bottom-dwelling or near-bottom prey. Those 
that have become restricted to feeding on food 
items buried within the bottom substratum tend 
to have a small, protrusible, inferior mouth and a 


strongly reinforced spade-shaped rostrum that 
aids in rooting out prey. The more generalized 
feeders tend to have a larger, more terminal 
mouth. Many of those that feed on fish, 
cephalopods, and swimming crustaceans are 
known to seek their prey well off bottom. 
Examination of stomach contents of the abyssal 
species Coryphaenoides armatus Hector, 1875, 
C. leptolepis Giinther, 1877, and C. filifer (Gil- 
bert, 1895) has revealed a high proportion of 


[233] 


234 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 7 


pelagic food items in their diet (Pearcy and Am- 
bler 1974; Haedrich and Henderson 1974). 
Dense aggregations of Coryphaenoides rupestris 
(Gunner, 1765) are known to rise regularly to 
100-300 m off bottom in the western North At- 
lantic (Pechenik and Troyanovskii 1970; Haed- 
rich 1974). Coryphaenoides acrolepis (Bean, 
1884), a species of the North Pacific, though 
primarily benthopelagic, has also been reported 
captured thousands of meters above bottom 
(Rass 1963:221; Makushok 1967:201; Iwamoto 
and Stein 1974:16). The bizarre, globose-headed 
species Squalogadus modificatus Gilbert and 
Hubbs, 1916, Macrouroides inflaticeps Smith 
and Radcliffe, 1912, and Echinomacrurus mollis 
Roule, 1916, are probably primarily bathy- 
pelagic, but they have been taken most often in 
bottom trawls. 

Most species of macrourids probably dwell at 
bathypelagic or mesopelagic depths as larvae or 
juveniles (Marshall 1965). A few species have 
adopted, as adults, an exclusively pelagic habit 
in the deep midwaters of the oceans. When he 
reviewed the bathypelagic macrourid fishes, 
Marshall (1964) considered only two species, 
Odontomacrurus murrayi Norman, 1939, and 
Cynomacrurus piriei Dollo, 1909, as _ being 
strictly confined to the midwaters of the ocean, 
although he suspected (and later, with Taning 
(1966), gave evidence) that Macrouroides in- 
flaticeps Smith and Radcliffe and Squalogadus 
modificatus Gilbert and Hubbs were probably 
primarily bathypelagic. O. murrayi is known 
mostly from the eastern Atlantic, although the 
holotype was taken in the Indian Ocean. C. 
piriei is known only from waters inside the Ant- 
arctic Convergence, where it abounds. 

In the present paper, we describe three new 
species of bathypelagic macrourids, all from the 
Pacific. Two are clearly referable to a tren- 
chantly distinct new genus, Mesobius. The third 
is referred to Nezumia, a genus of numerous, 
almost exclusively benthopelagic, species. The 
three new species are known only from captures 
made in midwater nets fished far off the bottom. 


METHODS 


Methods for taking measurements and counts 
generally follow procedures outlined by Hubbs 
and Lagler (1958), but are modified for mac- 
rourids as indicated by Gilbert and Hubbs (1916) 
and Iwamoto (1970). 

Proportional measurements are not given as 


the customary percentages of the standard 
length, because of the usual greater or lesser loss 
and abrupt regeneration of the caudal tip to form 
a pseudocaudal fin. In place of following Gilbert 
and Hubbs (1916) in using as a measurement 
base the length from snout tip to anus (which is 
highly variable due to the frequent distortion 
caused by the dorsal flexure of the head), it has 
become common to employ as the base the head 
length, from the extreme tip of the snout to the 
most posterior bony margin of the opercle. 

In the Material Examined sections, the 
museum catalog number for each lot is followed, 
in parentheses, by the number of specimens and 
their ranges of head length and total length. Lo- 
cality and capture data complete the information 
for each lot. Depths, when originally given in 
fathoms, have been converted to the nearest me- 
ter. Abbreviations generally follow those 
suggested in ‘“‘The Council of Biology, Editors 
Style Manual’’ (American Institute of Biological 
Sciences 1972). Others used imeludescen— 
cruise: HL—head length; IKMT—Isaacs-Kidd 
midwater trawl; m.w.o.—meters of wire out (of 
towing warp); R/V—research vessel; sta.— 
station; TL—total length. 

Museum abbreviations are as_ follows: 
CAS—California Academy of Sciences, San 
Francisco, California; FAKU—Faculty of Ag- 
riculture, Kyoto University, Maizuru, Japan; 
LACM—Museum of Natural History of Los 
Angeles County, California; PPSIO—P. P. Shir- 
shov Institute of Oceanology, Academy of Sci- 
ences USSR, Moscow; SIO—Scripps Institution 
of Oceanography, University of California, San 
Diego, La Jolla, California; USNM—National 
Museum of Natural History, Smithsonian In- 
stitution, Washington, D.C.; VIMS—Virginia 
Institute of Marine Science, Gloucester Pt., Vir- 
ginia; ZMUC—Zoological Museum, University 
of Copenhagen. 


ACKNOWLEDGMENTS 


We thank the many persons, who, in various 
ways, have helped us in the completion of this 
paper. The following provided or helped in the 
loan of specimens: E. H. Ahlstrom and H. G. 
Moser (National Marine Fisheries Service 
(NMEBS), La Jolla, California); M. A. Barnett 
(SIO); F. H. Berry (NMFS, Miami, Florida); T. 
M. Clarke (University of Hawaii); P. Four- 
manoir (ORSTOM, Noumea); K. Fujita (Tokyo 
University of Fisheries); A. Fleminger (SIO); L. 


HUBBS & IWAMOTO: NEW BATHYPELAGIC MACROURIDS 235 


N. Knapp (Smithsonian Oceanographic Sorting 
Center, Washington, D.C.); K. Kuronuma (To- 
kyo); R. J. Lavenberg (LACM); K. F. Liem and 
T. McLellan (Museum of Comparative Zoology, 
Harvard); D. F. Markle (VIMS); I. Nakamura 
(FAKU); J. Nielsen (ZMUC); N. V. Parin 
(PPSIO); R. H. Rosenblatt (SIO); and P. J. 
Struhsaker (NMFS, Honolulu). 

J. E. Fitch, California Department of Fish and 
Game, Long Beach, California, provided the 
slide used in Figure 3 along with information on 
the otoliths of Mesobius berryi. 

H. Geoffrey Moser very kindly and skillfully 
drew for us the early prejuvenile of Mesobius 
berryi (Fig. 6). Other illustrations were drawn by 
Ke PS Smith (Fig. 11); C. Pape (Figs. 2 & 4). 
(Other figures, 5, 7, 8, 10, were drawn by T. 
Iwamoto.) Radiographs were made by J. E. 
Gordon. 

For their critical review of the manuscript we 
thank William N. Eschmeyer, N. B. Marshall, 
and Richard H. Rosenblatt. 


Mesobius Hubbs and Iwamoto, new genus 


TyYPE-SPECIES.—Mesobius berryi Hubbs and 
Iwamoto. 

DrIAGNosis.—The two species of the genus are 
bathypelagic. Each has seven branchiostegal 
rays. The anus and the urogenital opening are 
surrounded by a raised margin of black naked 
skin and thus are removed from the origin of the 
anal fin by the width of that margin. A large light 
organ, lying anterior to the anus, abuts the an- 
terior wall of the rectum (Fig. 5). The anus, the 
pelvic fins, and anal-fin origin are displaced far 
forward on the short abdomen; the distance 
from the isthmus to the anal-fin origin slightly 
exceeds the interorbital width, and is less than 
two-thirds the postorbital length. The mental 
barbel is absent. The sensory pores of the head 
are small and inconspicuous. The dentition in 
both upper and lower jaws comprises narrow 
bands of small depressible teeth, with none en- 
larged; the upper-jaw tooth bands are widely 
separated at the symphysis. The body scales 
(Fig. 4A) are small and thin. Those near the dor- 
sal and ventral margins of the trunk and tail bear 
1-3 small, erect spinules on the exposed field; 
some of the scales along the flanks lack spinules. 
The scales on the head (Fig. 4B) are uniquely 
elongated; each bears I-3 rows of spinules that 
form low, sharp ridges with 2-9 spinules per 
row. The head and body are compressed. The 


eyes of adults are only moderately enlarged (or- 
bit diameter 23-31 percent of head length). The 
mouth is large (upper-jaw length 40-47 percent). 
The gill-membranes are narrowly attached to the 
isthmus and have a narrow posterior free fold. 
The wide opercular opening extends anteroven- 
trally to a vertical about one corneal diameter 
behind the orbit. The first (outer) gill-slit is re- 
stricted. The greatly reduced gas-bladder bears 
two retia and two gas glands. Some of the short 
caeca are bifid at the base. Postlarvae and pre- 
juveniles pass through a polka-dotted (“‘phalac- 
romacrurus’’) stage; adults are mostly black. 
Styloptery (referring to the stalked pectoral fins) 
of the postlarvae is extreme, reminiscent of the 
stylophthalmy (greatly stalked eyes) of the post- 
larvae of Idiacanthus. 

RELATIONSHIPS.—Mesobius is referable to 
that group of macrourine rattails that has in 
common the combination of seven branchio- 
stegal rays and the anus only slightly removed 
from the anal fin by the width of a broad, circum- 
anal margin of naked black skin. Related gen- 
era include Echinomacrurus Roule, Trachonurus 
Goode and Bean, Cetonurus Gunther, Para- 
cetonurus Marshall, and Sphagemacrurus Fow- 
ler. Of these genera, Mesobius seems closest 
to Echinomacrurus, a monotypic genus of the 
eastern North Atlantic and western Indian Ocean 
(Marshall 1973:600). 

These two genera are similar principally in 
having a relatively short trunk (with pelvic and 
anal fins and anus far forward), much reduced or 
lacking gas-bladder, black coloration, reduced 
or absent chin barbel, and small teeth in narrow 
bands in each jaw. These characters, however, 
may reflect convergent life styles rather than 
phylogenetic proximity. The degree to which 
each character is developed in Mesobius and 
Echinomacrurus as a_ specialization to a 
bathypelagic habit is generally the more extreme 
in Mesobius, a strictly bathypelagic fish, and 
less modified in Echinomacrurus, a sometime 
dweller of the bathypelagic realm. Interestingly, 
however, the gas-bladder is entirely lacking in 
Echinomacrurus (Marshall 1973:509), but is only 
regressed in Mesobius (a condition that one 
might not expect). In the two other bathypelagic 
genera, Cynomacrurus Dollo and Odontomac- 
rurus Norman, the gas-bladder is also regressed 
(Marshall 1964). Significantly, all of these 
bathypelagic genera have weakly ossified skele- 
tons. 


236 


Aside from the characters mentioned above 
(and until the internal morphology of both gen- 
era has been more closely compared), there ap- 
pear to be few characters that would indicate 
close relationship between Echinomacrurus and 
Mesobius. The differences between the two 
genera are more obvious. Whereas Mesobius 
has a compressed head with only a moderate 
development of mucous’ chambers, in 
Echinomacrurus the head is broad and inflated, 
‘due mainly to the very wide lateralis canals” 
(Marshall 1973:598). The mouth of Mesobius is 
large and almost terminal, whereas that of 
Echinomacrurus is small and inferior. The 
greatly elongate and closely seriated head scales 
of Mesobius contrast sharply with the small, 
round, non-imbricate scales of Echinomacrurus. 

Mesobius is far removed from three other 
strictly bathypelagic macrourids: Odontomac- 
rurus  murrayi, Cynomacrurus  piriei and 
Nezumia parini Hubbs and Iwamoto. Odon- 
tomacrurus and Cynomacrurus each has six 
branchiostegal rays, large fanglike, jaw teeth, 
and large open sensory pores on the head. 
Mesobius, in contrast, has seven branchiostegal 
rays, small teeth in a narrow band in the jaws, 
and no apparent open sensory pores on the 
head. Nezumia parini, like Mesobius, has seven 
branchiostegal rays and a light organ closely as- 
sociated with the anus, but the position of the 
periproct (distantly removed from anal fin and 
closer to pelvic fins), the presence of a strong 
suborbital ridge and a stout terminal snout scute, 
the relatively small, inferior mouth, and the re- 
stricted gill membranes, all indicate that 
Nezumia parini is a member of a phylogenetic 
line very divergent from that of Mesobius. It ap- 
pears then, that the bathypelagic macrourids 
have had a diverse origin, and that they are not 
from one evolutionary line but from several. 

DISTRIBUTION.—From our study material, the 
genus appears to be confined to the Pacific 
Ocean—M. berryi to the northeastern portion 
and M. antipodum to the southwestern portion. 
Dr. N. B. Marshall, however, writes (in litt., 17 
March 1976) of having found a young Mesobius 
in the DANA collection (sta. 1166 IV, 10°16’N, 
40°41'W), and that he now believes that the 
polka-dotted young reported by Backus ef al. 
(1965) as Sphagemacrurus? is also a Mesobius. 
He concludes: ‘‘I expect that Mesobius has a 
world-wide distribution between the subtropical 
convergences.”’ 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 7 


TABLE 1. FREQUENCY DISTRIBUTIONS OF SELECTED COUNTS 
OF Mesobius berryi HUBBS AND IWAMOTO. ABBREVIATIONS: 
First DorsAL FIN—1D.; PECTORAL FIN—1P.; PELVic FIN— 
2P; GILL-RAKERS ON MEDIAL Row OF First ARCH—GR I; 
GILL-RAKERS ON MEDIAL Row OF SECOND ARCH—GR II. 


7 8 J KD its i B18 a 
1D. 3 aes) 11.00 
lips Sy Ales 13.07 
ZR 1] ily 22 7.70 
GRI | 4.) 7, 3 11.80 
GR II PHY AWE 7) 11.00 


* Includes counts of both right and left fins from each 
specimen. 


EtTyMOLOGY.—The name Mesobius is derived 
from peaos (mesos), middle, and Bos (bios), 
life, referring to the bathypelagic habitat of the 
genus. 


Mesobius berryi Hubbs and Iwamoto, new 

species 

(FiGuRES 1-8, 10A; TABLES 1-3) 

““Macrouridae, unidentified’? Berry and Perkins 1966:668 
(‘‘an undescribed genus and species of bathypelagic mac- 
rouroid’’; two specimens from R/V Coss, cr. C6303, sta. 
86.92, 305 km SW of Point Conception, California, in ap- 
proximately 640 m.) 


DIAGNosIs.—A species of the genus Mesobius 
with 12-17 pyloric caeca; medial gill-rakers on 
first arch 10-13, on second arch 10-12; snout 
length 23-28 percent and interorbital width 
28-34 percent of head length. 

DESCRIPTION OF ADULTS.—Selected counts 


are given in Table | and measurements in Table 
2 


“=. 


General Adult Features: The general adult 
features of this unique fish may be seen in the 
illustration of the holotype (Fig. 1). The head 
and body are compressed; the greatest width of 
the head is about half the greatest body depth. 
The fin rays are fragile and slender, and the 
skeleton is poorly ossified. Most of the dermal 
head bones are thin, in part almost membranous. 
The integument of the head and body is thin, but 
is not membranous (as it is in members of the 
genus Hymenocephalus). The temporal region 
of the head is much enlarged. The high supraoc- 
cipital crest forms a distinct arch in the dorsal 
profile. Ventral aspects of the abdomen are 
short; the distance from the isthmus to the anal 
fin is about 1.0 1.5 orbit diameters. The anus, 
pelvic fins, and anal-fin origin are displaced 


237 


HUBBS & IWAMOTO: NEW BATHYPELAGIC MACROURIDS 


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238 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 7 


Ficure |. Mesobius berryi Hubbs and Iwamoto. Holotype, SIO 73-170, 61.8 mm HL, 344 mm TL, captured in midwaters of 


the central equatorial Pacific at an estimated depth of 0-1280 m. 


much farther forward than in most other mac- 
rourids (but about the same as in Odontomac- 
rurus murrayi and Cynomacrurus piriei and in 
species of Sphagemacrurus and of subgenus 
Lucigadus of genus Ventrifossa). The posterior 
margin of the opercular bones are finely scal- 
loped or crenulated. The interopercle is naked 
and scarcely exposed beyond the ventral and 
posterior margins of the large preopercle. The 
large lower jaw bears a pronounced knob at the 
symphysis. A small tubercle below the sym- 
physis may be the remnant of a chin barbel. The 
gills occupy a relatively small portion of the 
capacious gill cavity; the tubercular rakers are 
widely spaced along the gill-arches. The short 
gill-filaments, measuring less than 1.5 in pupil 
diameter, are poorly developed in comparison 
with those of benthopelagic relatives (Fig. 2). 
The pyloric caeca are short and thick (some are 
bifid at the base); four central Pacific specimens 
yield 12, 13, 16, and 17 tips. Eleven abdominal 
vertebrae were counted in five specimens. 

After examining four left-sagittal otoliths, 
John E. Fitch, California Department of Fish 
and Game, who provided the slide from which 
Figure 3 was taken, has stated (in litt., 14 July 
1975) that: 


‘‘Very few kinds of fish show such a radical change in 
otolith shape as does Mesobius. Typically lobular margins 
are a juvenile character, except among cods and a few 
others in which they are a family character. The three 
smallest Mesobius otoliths show differences other than 
the juvenile lobular margins.”’ 


Fins: The first dorsal is small, with the greatest 
height less than the postorbital length. The first 
of the two spinous anteriormost rays 1s small and 
thornlike; its sharp leading edge is armed in 
some specimens with one or two minute denti- 
cles. The second ray is much longer, more slen- 
der, and near the base it is triangular in cross- 
section; its rather sharp leading edge bears 
small, often irregular serrations in most small 
specimens and in some large ones. The low sec- 
ond dorsal is poorly developed throughout. The 
relatively slender, pointed, narrow-based pec- 
torals are inserted high on the body, much 
nearer the dorsal than the ventral body contour. 
The pelvics are small and slender. The anal is 
moderately well developed over its entire length 
and is much larger than the second dorsal. The 
pelvics originate below the posterior part of the 
preopercle; the anal origin lies below the pec- 
toral base and below the anterior third of the 
first dorsal. 

Scales: The scale structure is characterized in 
the diagnosis of the genus. The head is almost 
completely scaled except on the gill-membranes, 
lips, interopercle, and on the small narial fossa. 
In some specimens with heads longer than about 
40 mm, the body scales, except those ventrally 
on the abdomen, over the nape, and around the 
base of the first dorsal, are devoid of spinules. 
The scales on the pectoral base are very thin, 
non-spinulated, and highly deciduous (only a 
few remain on one specimen). The orientation of 


HUBBS & IWAMOTO: NEW BATHYPELAGIC MACROURIDS 239 


FIGURE 2. Right outer gill-arch of (A) Mesobius berryi and 
(B) Nezumia liolepis, comparing relative size of filaments of 
the two species. Drawn by Cherryl Pape. 


the elongate head scales (Fig. 4) and the as- 
sociated spinule rows render the head surface 
striated, in groups of varying, parallel courses. 
Over the temporal region the changing align- 
ments, in more or less sigmoidally curved 
groups, are particularly conspicuous, from the 
area behind the orbit upward and backward over 
the nape. 

Gas-bladder: The very small, elliptical blad- 
der has a small nib pointed posteriorly; in a 40- 
mm-HL specimen the organ is 6 mm long. The 
external tunica of the gas-bladder is thin and 
transparent, but tough. What appear to be two 
gas glands are enveloped in flocculent, white, 
lipoidal material. Retia, attached to the gas 
glands, emerge anteroventrally from the gas- 
bladder. 

Light Organ: Partial dissection of a 64-mm- 
HL paratype (SIO 72-374) indicates that this 
species has a relatively large oval or bean- 
shaped structure, presumably a light organ, en- 


Ficure 3. Otoliths (left sagittae) from four specimens of 
Mesobius berryi. Top to bottom: LACM 9585-20 (178 mm 
TL); LACM 30083-17 (26 mm HL, 190 mm TL); LACM 8638 
(33 mm HL, 170 mm TL); LACM 9579-16 (70 mm HL, 392 
mm TL). Otolith lengths are, respectively, 3.4 mm, 4.8 mm, 
5.3 mm, and 7.1 mm. Photograph provided by John E. Fitch. 


capsulated in a thin layer of muscle, and fronting 
the lower wall of the rectum. A midsaggital sec- 
tion through the organ (Fig. 5) reveals horizon- 
tally oriented folds of tissue with free borders 
posteriad. Anteroventral to the light organ is a 
small area, where the tissue is relatively translu- 
cent, that may serve as a ‘“‘window’’ through 
which light is transmitted to the exterior. The 
absence of a large internal lens fronting the light 


240 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 7 


A 


Ficure 4. Scales from Mesobius berryi: (A) from below first 
dorsal fin; (B) from posttemporal region of head. Drawn by 
Cherryl Pape. 


organ, so typical of most members of genera 
Malacocephalus, Ventrifossa, and Nezumia, in- 
dicates that Mesobius is a phylogenetically di- 
vergent member of the macrourid group charac- 
terized by having seven branchiostegal rays and 
an abdominal light organ. 

Coloration in Alcohol: Adults are black or 
brownish black over the entire head and body, 
with a violet tinge over the trunk region. The tail 
surfaces are somewhat paler than the more an- 
terior surfaces. 

DESCRIPTION OF YOUNG.—A postlarva of 12 
mm total length (SIO 72-14) has a head length of 
about 1.7 mm. The mouth is oriented almost ver- 
tically with the posterior end of the upper jaw 
lying on a vertical through the anterior edge of 
the eye. The diameter of the large eye measures 
less than 3 in HL, and is roundish, with a pos- 
terior bulge. The interorbital region is much nar- 
rower than the pupil. The abdomen is heavily 
pigmented with large melanophores. The anus is 
centrally located on the ventral midline. A black 
conical mass, the light organ, extends anteriad 
from the anus to between the bases of the pelvic 
fins. Rays of the anal fin appear to arise slightly 
anterior to the posterior end of the abdominal 
cavity. The pelvic-fin bases and the anus lie for- 
ward of the pectorals, which, in turn, lie slightly 
anterior to the origin of the first dorsal fin. The 
remarkably stylopterous pectorals are attached 
at the end of a slender pedicel about as long as 
the postorbital portion of the head. 


Rectum 


Light Organ ~ _ Genital Duct 


Pelvic Girdle __ Urinary Duct 


"Window" 


yO 
Pelvic Fin AN Anal Fin 


Sphincter Muscle \Sphincter Muscle 


Anus 


FiGureE 5. Diagrammatic illustration of midsaggital section 
through rectum and light organ of Mesobius berryi. 


Large prominent blotches occur on the snout, 
the tip of the lower jaws, the posterior angle of 
each side of the lower jaw, the posterior portion 
of the interorbital region, over the occipital re- 
gion of the skull, over the posttemporal region, 
on the preopercle just behind the orbits, and at 
the anterior edge of the base of the first dorsal 
fin. A series of uniformly spaced saddle marks 
extends along the dorsal margin of the tail, and 
dashlike marks along the ventral margin of the 
tail are spaced about half as far apart as are the 
dorsal saddle marks. 

An early prejuvenile (SIO 75-465; Fig. 6), 23.4 
mm in total length to tip of the broken tail, has 
essentially the same characteristics as those of 
the 12-mm specimen just described except that 
the periproct region, the anal origin, and the pel- 
vic fins are situated somewhat farther forward, 
and the dashlike marks on the ventral profile of 
the tail are larger and more widely spaced. The 
stylopterous pectorals are still well developed. 

In a 73-mm specimen (SIO 63-119), with head 
about 6.3 mm long, the mouth is still steeply 
oblique. The interorbital space is slightly nar- 
rower than the pupil. The anus and pelvic fin 
appear to have swung so far forward that they lie 
directly under the hind end of the opercle. The 
base of the pectoral fin is no longer pediculate 
but is reduced as in normal adults. Denticles 
along the leading edge of the second spinous 
dorsal ray are slender, sharp, and well devel- 
oped. Spots on the head and body closely re- 
semble those of a larger individual illustrated in 
Figure 7 (top). The saddles below the second 
dorsal and the dash marks along the anal are, 
however, more numerous and more prominent, 
especially posteriorly on the tail. Size-related 
changes in color pattern, from this size on, 
closely parallel those described for Odontomac- 
rurus murrayi by Marshall (1964). 

A prejuvenile specimen (Fig. 7, top) 15.7 mm 


HUBBS & IWAMOTO: NEW BATHYPELAGIC MACROURIDS 241 


FiGure 6. Early “‘phalacromacrurus”’ prejuvenile of Mesobius berryi, trawled in midwater at 31°15.0’N, 132°07.8’W, in the 
open eastern Pacific between southern California and Hawaii. Length 23.4 mm to broken tip of tail. Dorsal and anal rays 


diagrammatically shown. Drawn by H. Geoffrey Moser. 


in head length (LACM 35671), does not differ 
significantly in general features from individuals 
half as large. The medium-brown trunk becomes 
paler posteriad on the tail to tan. The head is tan 
with scattered punctulations, and the body and 
head display series of large, prominent, circular, 
brown blotches as illustrated. A small dark spot 
is located dorsomedially on the nape, and there 
is another on the tip of the lower jaw. (In a 
specimen of comparable size (SIO H51-87, 16.7 
mm HL), the spots are relatively smaller, and 
those along a file dorsally and midlaterally are 
more closely approximated.) The elongate 
scales on the head and the linear arrangement of 
the spinules on the scales are evident even at 
this early stage. Spinules are few on each scale, 
and they are slender, conical and erect. The cir- 
cular scales on the body generally have only one 
to two spinules, and some scales along the ven- 
tral margins of the body and tail completely lack 
spinules. 

In a somewhat larger individual (LACM 


30083-17) of about 26 mm HL (Fig. 7, bottom), 
the overall color is much darker, generally 
brownish black. The blotches of the body are 
relatively smaller and are becoming obliterated 
by the dark ground color; a spot along the pos- 
terior base of the first dorsal fin, however, is very 
dark. In head and body configurations, this stage 
already resembles that of the adult. The mouth is 
nearly horizontal, the interorbital region is broad 
and rounded, and the ventral aspects of the 
trunk have already shifted posteriad approxi- 
mately to their adult location. 

A still larger individual (SIO 73-329), 40 mm 
in head length, has essentially the adult config- 
uration and coloration—it is very blackish over 
the head and trunk, but slightly paler over the 
tail, on which faint traces of the lateral blotches 
still remain. 

COMPARISONS.—A comparison of specimens 
from the eastern and from the central Pacific 
discloses no significant regional differences. 
Selected measurements and counts are tabulated 


242 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 7 


FiGure 7. Diagrammatic illustrations of Mesobius berryi showing gross pigmentation patterns in juveniles 15.7 mm in head 
length (LACM 35671) (top) and 26 mm in head length (LACM 30083-17) (bottom). Fins and head partially reconstructed. Scales 


indicate 25 mm. 


(Table 3) for the comparison of specimens from 
these two regions, and for contrast with the 
single New Zealand specimen of Mesobius 
antipodum. The flabbiness of the fish and the 
consequent difficulty in making accurate mea- 
surements calls for some reservation in evaluat- 
ing the morphometric data. 

The peculiar polka-dotted pigmentation pat- 
tern in early-life stages of Mesobius is not 
restricted to the genus but is also characteristic 
of the young of Odontomacrurus , Malacoceph- 
alus , and, possibly, Sphagemacrurus. Maul and 
Koefoed (1950) described two polka-dotted 
macrourids as representatives of a new genus 
and species, Phalacromacrurus pantherinus , but 
Marshall (1964) showed that their specimens were 


juveniles of Odontomacrurus murrayi. Marshall 
(1964: Fig. 3) also figured a polka-dotted juvenile of 
Malacocephalus. Backus et al. (1965) illustrated 
what they assumed to be a juvenile Sphage- 
macrurus (taken in the eastern Atlantic) that hada 
pigmentation pattern almost identical with that of 
M. berryi. We have attempted to borrow that speci- 
men, which supposedly was deposited in the Mu- 
seum of Comparative Zoology, but the curators 
there have reported their inability to find it. 
Sphagemacrurus is not, as far as we know, 
known from the eastern Pacific. Marshall 
(1973:514) considered Macrurus trichiurus Gar- 
man and M. fragilis Garman as belonging to that 
genus, but the junior author’s examination of the 
holotypes has revealed that the former is clearly 


HUBBS & IWAMOTO: NEW BATHYPELAGIC MACROURIDS 243 


conspecific with Nezumia convergens (Garman) 
and the latter is probably close to Paracetonurus 
Marshall. The species of Sphagemacrurus have 
10-13 pelvic fin rays as compared with 9 or 
fewer in species of Mesobius. 

The genus Odontomacrurus is not known 
from the Pacific, as far as we know, although 
Fourmanoir (1969) reported two specimens of 
Phalacromacrurus pantherinus from _ the 
stomachs of lancetfish, Alepisaurus, taken off 
New Caledonia and New Hebrides. Mr. Four- 
manoir has informed us (in litt., 16 Dec. 1975) 
that the two specimens had apparently been dis- 
carded. 

Malacocephalus has not previously been 
known from the eastern Pacific, although repre- 
sentatives are found in the Hawaiian Islands, off 
Japan, in the Philippines, and in Australian wa- 
ters. We have, however, examined one adult 
specimen of a Malacocephalus (Santa Barbara 
Natural History Museum no. 0061) taken off 
Santa Barbara, California. It represents the only 
record of the genus from the eastern Pacific. The 
polka-dotted juveniles of Malacocephalus are 
readily distinguishable from those of Mesobius 
by having a chin barbel (developed even in a 
specimen (VIMS 03922) of Malacocephalus sp. 
8 mm in HL), a pale esophageal portion of the 
stomach (blackish in Mesobius), and a promi- 
nent scaleless area anterior to the periproct. 

DISTRIBUTION.—Mesobius berryi has been 
captured in midwater nets fished as shallow as 
0-313 mm to as deep as (+2700 m. Most of the 
captures have been in trawls hauled to at least 
650-1000 m, in waters off southern California, 
off Baja California (Mexico), off Oahu in the 
Hawaiian Islands, and in the central North 
Pacific 8-10 degrees of latitude north of the 
Hawaiian Islands, and in the equatorial mid- 
Pacific off Christmas Island (Fig. 8). The species 
appears to have a distribution pattern similar to 
that of the myctophiform fish, Evermannella 
ahlstromi, which is apparently ‘‘limited to areas 
transitional in properties between major Pacific 
upper water masses’’ (Johnson and Glodek 
1975:725). Unlike E. ahlstromi, however, 
Mesobius berryi does not extend into the east- 
ernmost sector of the Pacific equatorial water 
mass but is apparently replaced there by 
Nezumia_ parini, the only other known 
bathypelagic member of the family in the eastern 
Pacific Ocean. 


ETYMOLOGY.—The species is named for 


TABLE 3. Mesobius spp. COUNTS AND MEASUREMENTS 
COMPARED BY LOCALITY. PROPORTIONAL MEASUREMENTS 
IN PERCENTAGE OF HEAD LENGTH. MORPHOMETRIC DATA 
FOR SPECIMENS SMALLER THAN 33 MM HL EXCLUDED. 


anti- 


Species berryi podum 
Location Eastern Central New 
Pacific Pacific Zealand 

No. of specimens 4 8 1 
COUNTS: 
Gill-rakers on 

first arch 11—13* 10-13 15 
Gill-rakers on 

second arch 10-12* 10-12 16 
Pyloric caeca 12-17} ca. 38 
MEASUREMENTS: 
Total length (mm) 170-392 257-397 390 
Head length (mm) 33-70 49-65 75 
Snout length 24-27 26-28 32 
Interorbital 

length 30-34 28-30 36 
Preanal length 99-103 98-102 114 
Outer pelvic ray 

to anal origin 12-13 9-14 19 
Isthmus to anal 

origin 27-33 25-33 40 
Body depth 77-719 74-81 84 


* Counts from seven specimens. 
+ Counts from four specimens. 


Frederick H. Berry, now of the Southeast 
Fisheries Center, Miami Laboratory, of the Na- 
tional Marine Fisheries Service, who first col- 
lected the species, brought it to the attention of 
the senior author, and reported it as ‘*Mac- 
rouridae, unidentified’ (Berry and Perkins 


1966:668). 

MATERIAL EXAMINED.—Holotype. SIO 73-170 (61.8 mm 
HL, 344 mm TL): mid-equatorial Pacific, 00°03.8’-07.0'N, 
154°56.0’W, midwater trawl; estimated fishing depth 0-1280 
m, 3000 m.w.o.; R/V GEORGE MELVILLE cr. CATO 2, tow 82, 
17 July 1973. 

Paratypes. Eastern Pacific (8 specimens).—LACM 35671 (1, 
15.7 mm HL, 165 mm TL): Mexico, off Isla Guadalupe, 
29°30.0'N, 119°10’W; IKMT; 0-313 m, 3000 m.w.o., bottom depth 
1850 m; R/V VELERO sta. 9904, 8 Aug. 1964.—LACM 8638 (1, 
about 33 HL, 170 TL); California, Santa Catalina Basin, 
33°22'15"N, 118°45'00”W; midwater trawl; 1313-1298 m, 7000 
m.w.o.; R/V VELERO sta. 10259, 13 Jan. 1965—LACM 
9579-16 (1, 70 HL, 392 TL): Mexico, off Isla Guadalupe, 
30°30'20"N, 118°49'59"W; IkMT; about 1273 m.w.o.; R/V 
VELERO Sta. 11226 30 Aug. 1966.—LACM 30083-17 (1, about 
26 HL, 190 TL): Mexico, off Isla Guadalupe, 28°15'00’N, 
118°12'00"W; 10-ft 1kKMT; about 919 m.w.o.; R/V VELERO Sta. 
12079, 14 April 1968.—SIO H51-87 (1, 16.7 HL, 182 TL): 


244 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 7 


208 


Ficure 8. Distributions of Mesobius berryi (circles), M. antipodum (squares), and Nezumia parini (triangles). 


Mexico, off Baja California Norte, 25°29’-31'N, 115°24’- 
09’'W; midwater trawl; 1902 m, 21 Mar. 1951.—SIO 63-110 (1, 
about 6.3 HL, 73 TL): eastern Pacific W of Mexico, 27°22.6'N, 
125°20.0'W; IKMT; Q-about 2700 m, 4500 m.w.o.; R/V 
Horizon cr. 6204H, sta. 100.160, 16 Apr. 1962.—USNM 
215327, formerly SIO 63-374 (2, 66-69 HL, 333-374 TL): 
California, WSW of San Diego, 32°07.3’-00.5'N, 122°39.5’— 
41.4’W; Cobb Pelagic Trawl Mark II; fishing depth about 640 
m, bottom depth 4209 m, 1738 m.w.o.; R/V Coss cr. 58-7, sta. 
86.92, 6 Mar. 1963. 

Central Pacific (10 specimens).—CAS 20470 (1, 62 HL, 316 
TL): SW of island of Oahu, Hawaii, 21°20’-30'’N, 158°20’— 
30'W; 10-ft IkKMT; fishing depth 1150 m, 3300 m.w.o.; R/V 
TERITU sta. 70-920, 21 Sept. 1970.—CAS 30469 (1, 49 HL, 
257 TL): data as previous lot except depth about 1000 m, 
2850-3000 m.w.o.; sta. 70-12-13, 11-12 Dec. 1970.—CAS 
30471 (1, 61 HL, 340 TL): data as previous lot except depth 
950-1000 m, 2700 m.w.o.; sta. 70-12-30, 16 Dec. 1970.— 


USNM 215326 (2, 57-62 HL, 307-345 TL): data as previous lot 
except depth 700-800 m, 2000 m.w.o.; sta. 739-24, 25 Sept. 
1973.—SIO 72-14 (1, about 1.7 HL, 120 TL): 27°20.6’—23.5'N, 
155°20.7'-23.5'W, 1k plankton trawl, 1000 m.w.o.; R/V 
THOMAS WASHINGTON Exped. Aries 9, sta. PSI, 26 Sept. 
1971.—SIO 72-374 (1, 64 HL, 358 TL): 30°56.7’-59.7'N, 
155°24.3'-16.3'W; Isaacs-Brown closing midwater trawl; fish- 
ing depth 970 m, 1700 m.w.o.; R/V GEORGE MELVILLE cr. 
CATO I, tow 41, 30 June 1972.—SIO 73-329 (1, about 40 HL, 
221 TL): 28°03.5'-12.9'N, 154°38.8’-36.5'W; 10-ft IKMT; 
3000 m.w.o.; R/V THOMAS WASHINGTON cr. TASADAY I, tow 
42, 22 June 1973.—SIO 73-327 (1, 65 HL, 397 TL): 27°58.6’— 
28°07.2'N, 154°57.8’-155°07.5'W; 10-ft IKMT; 3000 m.w.o.; 
R/V THOMAS WASHINGTON cr. TASADAY I, tow 17, 18 June 
1973.—SIO 75-465 (1, 23.4 mm TL): 31°15.0’N, 
132°07.8'W; oblique plankton tow hauled obliquely between 0 
and 141 m, bottom depth 4691 m; R/V Horizon cr. H6204-05, 
sta. 70.200, 31 Mar. 1962. 


HUBBS & IWAMOTO: NEW BATHYPELAGIC MACROURIDS 245 


Ficure 9. Holotype of Mesobius antipodum Hubbs and Iwamoto. FAKU 47812, about 75 mm HL, 390 mm TL with tail 


incomplete, from off South Island, New Zealand, in 995-1110 m. 


Mesobius antipodum Hubbs and Iwamoto, new 
species 
(FIGURES 8, 9, 10B; TABLE 3.) 


DIAGNOSIS (based on one specimen).—A 
species of the genus Mesobius with about 38 
small, branched pyloric caeca; medial gill-rakers 
on first arch 15, on second arch 16; snout length 
about 32 percent of head length; interorbital 
width about 36 percent of head length. 

Counts.—First dorsal fin 11,9; pectoral fin 
14; pelvic fin 7/7; gill-rakers in medial row 
2 + 13 on first arch and 2 + 1 + 13 on second 
arch; scales below origin of first dorsal fin about 
13-14; scales below middle of first dorsal fin 
about 8-9. 

MEASUREMENTS.— Total length 390 mm (with 
an incomplete tail); head length about 75 mm. 
The following proportional measurements are in 
thousandths of head length: postrostral length of 
head 720; snout length 320; orbit diameter 227; 
interorbital width 360; postorbital length of head 
480; orbit to angle of preopercle 467; suborbital 
width 160; upper-jaw length 427; preanal length 
1133; outer pelvic ray to anal origin 187; isthmus 
to anal origin 400; greatest body depth 840; in- 


terspace between first and second dorsal fins 
533; length pelvic fin 267; length outer gill-slit 
12? 

DESCRIPTION AND COMPARISONS.—Most fea- 
tures of this fish closely parallel those described 
for its congeneric relative Mesobius berryi. The 
high pyloric caeca count in this species results 
not from a larger caecal mass than found in ber- 
ryi but rather from a greater branching of the 
mass to form more, and smaller, individual 
caeca. Larvae and juveniles are unknown, but 
we presume that individuals pass through a 
polka-dotted “‘phalacromacrurus”’ stage, as in 
M. berryi. 

Because of the relatively poor condition of the 
holotype. and only specimen, precise mea- 
surements could not be made of most features. 
Differences between M. antipodum and M. ber- 
ryi in morphometry (Table 3) may prove insignif- 
icant when additional specimens of antipodum 
are studied. Other features (meristic ones in par- 
ticular), however, seem to confirm the differ- 
ences suggested by the morphometric data. The 
holotype of M. antipodum appears to represent 
a much stouter, heavier-bodied species: the 
greatest breadth of the trunk is approximately 


246 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 7 


B 


Figure 10. Diagram of posterior region of head of (A) 
Mesobius berryi and (B) M. antipodum, comparing the pos- 
terior extent of the posttemporal region (solid arrows) and the 
relative curvature of the subopercular margin (open arrows). 


half the postrostral length of the head, whereas 
in M. berryi it is less than one-third that dimen- 
sion. Although the holotype of M. antipodum, at 
75 mm HL, is larger than the largest available 
specimen of M. berryi, the posterior nostril, at 
2.8 mm, is much smaller (3.7 mm) than it is in the 
largest M. berryi specimen (LACM 9579-16, 70 
mm HL). The shape of the opercle and subor- 
percle in the two species appears to differ (Fig. 
10; cf. also Figs. | and 9). The posterodorsal arm 
of the opercle is relatively much longer in berryi 
than in antipodum, and the posterior margin of 
the subopercle is more strongly incurved in an- 
tipodum. Finally, the sharply marked posterior 
margin of the posttemporal region (Fig. 10) in 
this species lies above the base of the pectoral 
fin, whereas in M. berryi it scarcely extends be- 
yond the posterior margin of the opercle. 


Because only one specimen of M. antipodum 
is known, little can be said about the distribution 
of the species. It would not be improbable to 
find that the species is widely distributed across 
the southern hemisphere, especially in the east- 
ern sector, in temperate to tropical waters. 

ETYMOLOGY.—The name is derived from the 
Latin antipodum, genitive of ‘‘antipodes,’’ re- 
ferring to persons dwelling at opposite points on 


the globe. 

MATERIAL EXAMINED.—Holotype. FAKU 47812 (about 75 
mm HL, 390 mm TL with tip of tail broken off): New Zealand, E 
of South Island, 44°44’S, 175°42.0’E; 995-1110 m; KAlyo MARU 
sta. 36, 16 July 1968. 


Nezumia parini Hubbs and Iwamoto, new 
species 
(FIGURES 8, 11; TABLEs 4, 5.) 


“‘A new bathypelagic species of Nezumia,’”’ Parin, Becker, 
Borodulina and Tchuvassov, 1973 (from English summary, 
p. 159). 

Nezumia sp.: Parin, 1975: 321 (2 specimens from W of 
Galapagos). Parin, Pokhilskaya, Sazonov and Fedoryako, 
1976: 225 (description; 2 specimens). 


DiAGNosis.—This_ distinctive bathypelagic 
species of Nezumia is black. Each of the small, 
circular scales bears I-15 long, slender, erect 
spinules that render the entire body surface vel- 
vety. Scale rows below the origin of the second 
dorsal fin number about 11-14, and there are 
about 50 scales in the lateral line from its origin 
Over a space equal to the predorsal length. The 
outermost of the short gill-filaments are half as 
long as the pupil. The gill-rakers on the inner 
side of each of the first two gill-arches number 
10-13 (usually 11-12). The outermost of the 
11-12 pelvic rays is usually greatly prolonged, 
1.0 to 2.5 times as long as the head. The gas- 
bladder is rudimentary. The weakly developed 
lateral line is represented by a file of small papil- 
lae. 

DESCRIPTION.—General Features: Counts 
and measurements are given in Tables 4 and 5. 
The greatest width of the moderately compres- 
sed head is about two-thirds the greatest body 
depth. The bluntly pointed snout is armed with a 
spiny median scute and small, blunt, lateral 
scutes. The interorbital region is gently convex. 
Except in young individuals, the orbit does not 
quite enter the dorsal profile. The gill- 
membranes are broadly attached to the isthmus. 
The outer gill-slit is restricted. The small, tuber- 
culear gill-rakers number 1-3 + 910 (total 


HUBBS & IWAMOTO: NEW BATHYPELAGIC MACROURIDS 247 


Ficure 11. Holotype of Nezumia parini Hubbs and Iwamoto, SIO 73-165, 34 mm HL, 210 mm TL, from central equatorial 
Pacific. Enlarged dorsal and lateral views of scale from region between origin of second dorsal fin and lateral line (not drawn to 


scale). Scale indicates 25 mm. Drawn by Katherine P. Smith. 


10-13) on the mesial side of the first arch, and 
1-2 + 8-10 (total 10-12) on the second arch. The 
gill-filaments are much shorter than those in the 
most closely related benthopelagic relatives and 
appear to be more fragile. The anus is located 
within the middle third of the distance between 
the pelvic-fin base and the anal-fin origin. The 
anterior dermal window of the midventral light 
organ is externally evident as a small, teardrop- 
shaped fossa adjacent to the anus. The relatively 
short, thin, simple pyloric caeca branch near 
their attachment to the wall of the pylorus; 21-29 
caeca were counted in four specimens. In a male 
paratype (SIO 73-165) 30 mm in head length, the 
deflated rudimentary gas-bladder is only 2.5 mm 
long. Its drumming muscles are large (about 6 
mm long), but have no apparent direct connec- 
tion with the gas-bladder; they originate an- 
teriorly on the body wall and are probably in- 
serted posteriorly to the mesentary around the 
gas-bladder. The lateral line is reduced to a 
series of small, black papillae (free neuromasts?) 


that follow a course, from the posttemporal re- 
gion posteriad onto the trunk and tail, similar to 
that of the lateral line in most macrourids. The 
sensory pores on the head are poorly developed. 

Scales: The small, almost circular scales are 
armed with I-15 (more in large specimens) long, 
slender, erect, terminally recurved spinules, ar- 
ranged in |-4 slightly divergent rows. Almost 
the entire head and body are uniformly scaled 
except on the ventral and anterodorsal surfaces 
of the snout, portions of the suborbital region, 
the gill-membranes, and the periproct. The base 
of the first dorsal, pelvic, and pectoral fins, and 
the mandibular rami, are sparsely and loosely 
scaled. The margins of the mandibles are often 
devoid of scales. 

Fins: The fins are moderately well developed, 
although the rays are fragile. The outer pelvic 
ray is prolonged into a hairlike filament. The 
pelvic fins lie directly below the pectorals and 
anterior to the dorsal (these relative fin positions 
are somewhat exaggerated in the holotype (Fig. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 7 


248 


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HUBBS & IWAMOTO: NEW BATHYPELAGIC MACROURIDS 


TABLE 5. RANGE, MEAN (X) AND STANDARD DEVIATION 
(S.D.) oF SELECTED COUNTS TAKEN FROM TyPE-SPECIMENS 
oF Nezumia parini HUBBS AND IWAMOTO. ABBREVIATIONS: 
First DorsAL Fin—1D.; PECTORAL FIN—1P.; PELvic FiInN— 
2P.; GILL-RAKERS ON MEDIAL SIDE OF FIRST ARCH, IN- 
CLUDING RUDIMENTS—GR I; GILL-RAKERS ON MEDIAL SIDE 
OF SECOND ARCH—GR II. 


N Range x Sy By. 
1D. 20 9-11 10.30 0.80 
ips 36 13-19 16.03 1.30 
2p 39 10-12 11.00 0.32 
GR I 20 10-13 11.40 0.68 
GR Il 20 10-14 LESS 0.89 


* Includes counts of both right and left fins from each 
specimen, if possible. 


11) due to some dorsal flexure of the head in 
preservation). The anal fin originates slightly 
behind a vertical through the end of the first dor- 
sal base. The first dorsal spine is small and 
spikelike; the second is slightly prolonged and is 
armed along the leading edge with a file of small 
sharp spinelets. 

Dentition: The small teeth form narrow bands 
in each jaw. Those on the premaxillary are nar- 
rowly separated at the symphysis, and those of 
the outer series are slightly enlarged. 

Coloration in Alcohol: Some specimens are 
almost entirely black; others grade to dark 
brownish black. The buccal cavity is dark gray, 
but the gill cavity and the gill-filaments are 
mostly pallid. The peritoneal membrane is a 
splotchy dark brown. 

DISTRIBUTION.—Nezumia parini has been 
taken only in open midwater nets fished at meso- 
to bathypelagic depths in the eastern and 
equatorial central Pacific (Fig. 8). The species is 
found in a narrow belt along the equator, from 
the mid-Pacific to the coast of South America. 
Its distribution along the continental margin ap- 
pears to stop at the Gulf of Panama in the north, 
and northern Chile (at about 20°S) in the south. 
The northern boundary of its range coincides 
with the southern extent of the oxygen-minimum 
zone off Central America (Brandhorst 1959: Fig. 
4). Whether this zone is limiting to the dispersal 
of N. parini, as it is with many other midwater 
fishes, may seem doubtful in view of the species’ 
abundance off Peru, where another oxygen- 
minimum layer is developed. Perhaps the 
greater thickness of the layer off Central 
America, where it may be more than 1200 m 


249 


thick, forms an effective barrier to the north- 
ward dispersal of N. parini, while the less thick 
layer off Peru (maximum thickness about 800 m 


fide Wyrtki 1967) allows the species to reside 


below the oxygen depleted layer. 

RELATIONSHIPS.—In its evolution Nezumia 
parini appears to have undergone considerable 
selection rendering it better adapted to a 
bathypelagic existence. These changes, how- 
ever, have not obscured its basic relationship 
with that group of strictly benthopelagic species 
of Nezumia including N. aequalis (Gunther) and 
N. condylura Jordan and Gilbert. Characters 
that indicate this relationship include: (1) the 
relatively sharply pointed, broad rostrum armed 
with a large spiny terminal and two lateral 
scutes; (2) the markedly angular and prominent 
suborbital ridge; (3) the relatively small and in- 
ferior mouth; (4) the restricted openings be- 
tween the gill-arches; (5) the broadly connected 
gill-membranes; (6) and the relatively slender 
body. In varying degrees these characters con- 
trast with those of the generally much larger 
species, that have in common a higher, blunter, 
narrower snout, a flatter suborbital region, a 
larger and more terminal mouth, wider openings 
between the gill-arches, less restricted gill- 
membranes, and a more laterally compressed 
and deeper body and head. Those contrasting 
species include Nezumia atlantica (Parr, 1946), 
N. africana (Iwamoto, 1970), N. stelgidolepis 
(Gilbert, 1891), N. liolepis (Gilbert. 1891), N. 
holocentrus (Gilbert and Cramer, 1897), N. pu- 
dens Gilbert and Thompson, 1916, and N. 
bubonis Iwamoto, 1974. 

EVOLUTIONARY REMARKS.—The discovery of 
a bathypelagic species of Nezumia is remarkable, 
considering that its closest relations are with 
species that are specialized toward feeding on 
benthic organisms rooted from the bottom. That 
Nezumia parini, in evolutionary terms, has not 
been long removed from a benthopelagic habit is 
reflected in its retention of features prevailing in 
its nearest relatives: in particular, a stout bony 
rostrum and suborbital ridge, a mental barbel. 
and a protrusible, inferior mouth. N. parini ap- 
pears to have evolved the following features in 
adaptation to conditions in the bathypelagic 
realm: (1) entirely black or blackish color: (2) 
rudimentary gas-bladder; (3) short and thin gill- 
filaments: (4) vestigial lateral line: (5) small 
scales and thin, erect spinules that render the 
surface velvety: (6) much gelatinous intermuscu- 


250 


lar tissue; and (7) weak fin rays. Most of these 
features are shared in common with other 
strictly bathypelagic macrourids, as discussed 
by Marshall (1964). One can readily appreciate 
the advantages conferred by black color 
(camouflage), vestigial gas-bladder, reduced 
gill-filaments, gelatinous tissue, and weak fin 
rays (related to energy conservation and a 
slower life style). The peculiar scales are a puz- 
zle, however—what advantages would they 
give? This scale type is developed in Odon- 
tomacrurus, Cynomacrurus, Squalogadus, 
Macrouroides, and, to varying degrees, in 
Mesobius and Echinomacrurus. In contrast to 
the small eyes found in these other genera (ex- 
cepting Mesobius, which has moderate-size 
eyes), the eyes of N. parini are huge, measuring 
a third to two fifths of the head length. Perhaps 
the eyes are used more for detection of predator 
or prey than are the organs of the lateralis sys- 
tem (which seem to be less developed than those 
of its benthic relatives, especially along the 
trunk and tail). Feeding habits of N. parini are 
undoubtedly quite different from those of Odon- 
tomacrurus murrayi and Cynomacrurus piriei, if 
the large fanglike teeth, large terminal mouth 
and large, open head pores of the sensory 
lateralis system in the last two species are any 
indication. Feeding habits of the new species 
probably more closely approach those of 
Squalogadus modificatus and Macrouroides in- 
flaticeps, both of which have small, protrusible, 
inferior mouths that are probably adapted to 
feed on small pelagic invertebrates and fishes. 
Juvenile macrourids taken at bathypelagic 
depths are often crammed with bathypelagic 
copepods. Perhaps N. parini feeds on similar or- 
ganisms. 

ETYMOLOGY.—We take deep pleasure in nam- 
ing this highly distinctive species for our es- 
teemed Soviet colleague Nikolay V. Parin, who 
has also recognized it as undescribed, and has 
very generously made his material available to 


US. 

MATERIAL EXAMINED.—Holotype. SIO 73-165 (34 mm HL, 
210 mm _ TL): mid-equatorial Pacific, 00°01.3’-0.2’'S, 
155°O1.1’-154°59.8'W; 1KMT; oblique tow, 1500 m.w.o.; R/V 
GEORGE MELVILLE, cr. CATO 2, sta. 36, tow 71, 15 July 1972. 

Paratypes. Gulf of Panama: ZMUC P372830 (1, 14.8 HL, 82 
TL); 7°30'N, 79°19'W; stramin net with 150-cm opening; esti- 
mated fishing depth 0-1250 m, 2500 m.w.o., bottom depth 
2550 m; DANA sta. 1203 XII, 11 Jan. 1922 —ZMUC P372831 
(1, 17.7 HL, 90 TL); 6°48'N, 80°33’W; 300-cm ring-trawl; es- 
timated fishing depth 0-1750 m, 3500 m.w.o.; DANA sta. 1208 
IV, 16 Jan. 1922. Galapagos: PPSIO (1, 31 HL, + 164 TL); 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 7 


00°17.5'S, 89°28.7'W; IKMT; 0-600-700 m; R/V AKADEMIK 
KURCHATOV sta. 313, sample N261, 10-11 Nov. 1968. 
Ecuador: PPSIO (1, 32 HL, 198 TL); 00°00.0’'N, 85°00.0'W; 
IKMT; 1500-0 m; R/V AKADEMIK KURCHATOV sta. 219A, 
sample N25, 28 Aug. 1968. Peru: PPSIO (1, 18 HL, + 81 TL); 
5°54.8'S, 84°53.8'W; IKMT; 0-1400 m; R/V AKADEMIK 
KURCHATOV sta. 227, sample N65, 5 Sept. 1968 —USNM 
215325 (1, 19 HL, + 108 TL); 7°43’-33'S, 80°43'W; IKMT; 
700-1110 m; R/V ANTON BRUUN cr. 18B, sta. 755, 6 Sept. 
1966.—LACM 10279 (1, 18 HL, 90 TL); 7°47.5'S, 81°23'W; 
IKMT; 677 m; ELTANIN sta. 34, 7 June 1962.—CAS 29194 (1, 
20.5 HL, 118 TL); 8°26’S, 80°26’W; 1kmMT; 2150-1230 m; R/V 
ANTON BRUUN cr. 16, sta. 650H, 9 June 1966.—CAS 15996 (3, 
17-24 HL, 111-135 TL); 11°50’—52'S, 78°16’—-24'W; IKMT; 
0-1350 m, bottom depth 1640 m; R/V ANTON BruuwuN cr. 16, 
sta. 656P, 16 June 1966.—SIO 65-603 (1, 33 HL, 161 TL); 
11°51’-12°06'S, 79°04'-78°58.5'W; 10-ft IKMT; ca. 0-2000 m, 
5630 m.w.o.; R/V ANTON BRUUN cr. 12, sta. 1, 21 Nov. 
1965.—CAS 29414 (1, 35 HL, 216 TL); 11°52’-53'S, 78°24’— 
19’W; 1KMT; 0-900 m; R/V ANTON BRUUN cr. 16, sta. 656Q, 
13 June 1966.—USNM 215324 (1, 16.8 HL, 104 TL); 11°56’— 
55'S, 79°06'—78°53'W; IKMT; 0-455 m, bottom depth 4450 m; 
R/V ANTON BRUUN cr. 16, sta. 656A, 12 June 1966.—PPSIO 
(2, 26-37 HL, 140-221 TL); 12°20.0'S, 81°42.5’W; IkmT; 
0-1500 m; R/V AKADEMIK KURCHATOY sta. 229, sample N80, 
7-8 Sept. 1968.—PPSIO (1, 23 HL, + 125 TL); 12°43.9’S, 
78°31.9'W; IKMT; 0-1500 m; R/V AKADEMIK KURCHATOV Sta. 
277, sample N198, 25 Oct. 1968.—SIO 72-195 (4, 15-32 HL, 
75-192 TL); 13°51.3’S, 77°41.1'W; 10-ft IkKMT; estimated fish- 
ing depth 0-1100 m, 3000 m.w.o.; R/V WASHINGTON South- 
Tow Exped., sta. MV72-II-38, 12-13 May 1972. Chile: SIO 
72-182 (1, 21 HL, 90 TL); 18°38.4'-35.1’S, 70°39.0’-35.1'W; 
10-ft IKMT; estimated fishing depth 0-940 m, 2500 m.w.o.; 
R/V WASHINGTON SouthTow Exped., sta. MV72-II-25, 6-7 
May 1972.—PPSIO (1, 35 HL, 230 TL); 20°00.0'S, 76°42.0'W; 
IKMT; 0-1500 m; R/V AKADEMIK KURCHATOV Sta. 236, sam- 
ple N112, 17 Sept. 1968. Equatorial Pacific: PPSIO (1, 16 HL, 
+ 104 TL); 00°01.4’S, 97°02.7'W; 1kmT; 0-700 m; R/V 
AKADEMIK KURCHATOV sta. 1454, sample N98, 18 Jan 
1974.—PPSIO (1, 14 HL, + 70 TL); 00°01.4’S, 97°02.7'W; 
IKMT; 0-410-420 m; R/V AKADEMIK KURCHATOV sta. 1454, 
sample N112, 19 Jan. 1974.—SIO 69-496 (1, 29 HL, 142 TL); 
02°20'-12'S, 100°42.5'-27'W; 10-ft IKMT; 0-675 m, 1800 
m.w.o., bottom depth 3292 m; R/V WASHINGTON sta. QBR— 
16-MWT, 26 Nov. 1969.—SIO H52-38 (2, 19-22 HL, 118-154 
TL); 00°17.7'N, 110°26’W; 7 June 1952.—PPSIO (1, 38 
HL, 245 TL); 01°59’N, 125°19’W; 1kMT; 0-900-1000 m; R/V 
LIRA trawl 51, 27 Mar. 1966.—SIO 73-171 (1, 36 HL, 228 TL); 
00°07.0'-06.6'S, 154°56.0'-155°01.4'W; IKMT; oblique tow, 
3000 m.w.o.; R/V MELVILLE cr. CATO 2, sta. 36, tow 83, 17 
July 1972.—SIO 73-165 (1, about 30 HL, 185 TL); data same 
as for holotype. 


LITERATURE CITED 


BACKUS, RICHARD H., Gites W. MEAD, RICHARD L. HAED- 
RICH, AND ALFRED W. EBELING. 1965. The mesopelagic 
fishes collected during cruise 17 of the R/V CHAIN, with a 
method for analyzing faunal transects. Bull. Mus. Comp. 
Zool. 134(5): 139-158. 

BERRY, FREDERICK H., AND HERBERT C. PERKINS. 1966. 
Survey of pelagic fishes of the California Current area. 
U.S. Fish Wildl. Serv., Fish. Bull. 65(3): 625-680. 

BRANDHORST, WILHELM. 1959. Nitrification and denitrifica- 
tion in the eastern tropical North Pacific. J. Cons. Perm. 
Int. Explor. Mer. 25: 3-20. 


HUBBS & IWAMOTO: NEW BATHYPELAGIC MACROURIDS 


FouRMANOIR, P. 1969. Contenus stomacaux d’Alepisaurus 


(Poissons) dans le  Sud-Ouest Pacifique. Cah. 
O.R.S.T.O.M., sér. Oceanogr. 7(4): 51-60. 
GILBERT, CHARLES H., AND CARL L. Husss. 1916. Report 


on the Japanese macrouroid fishes collected by the United 
States fisheries steamer *‘Albatross”’ in 1906, with a synop- 
sis of the genera. Proc. U.S. Natl. Mus. 51(2149): 135-214, 
pls. 8-11. 

HAEDRICH, RICHARD L. 1974. Pelagic capture of the epiben- 
thic rattail Coryphaenoides rupestris. Deep-Sea Res. 21: 
977-979. 

, AND N. R. HENDERSON. 1974. Pelagic feeding by 
Coryphaenoides armatus, a deep benthic rattail. Deep-Sea 
Res. 21: 739-744. 

Husss, Cart L., AND KARL F. LAGLER. 1958. Fishes of the 
Great Lakes region. (Rev. ed.) Cranbrook Inst. Sci. Bull. 
26: 1-213. 

Iwamoto, Tomro. 1970. The R/v PiLLsBuRY Deep-Sea 
Biological Expedition to the Gulf of Guinea, 1964-65. 19. 
Macrourid fishes of the Gulf of Guinea. Stud. Trop. 
Oceanogr. Miami 4(pt. 2): 316-431. 

, AND Davip L. STEIN. 1974. A systematic review of 
the rattail fishes (Macrouridae: Gadiformes) from Oregon 
and adjacent waters. Occas. Pap. Calif. Acad. Sci. 111: 
1-79. 

JOHNSON, ROBERT K., AND GARRETT S. GLODEK. 1975. 
Two new species of Evermannella from the Pacific Ocean, 
with notes on other midwater species endemic to the Pacific 
central or the Pacific equatorial water masses. Copeia 
1975(4): 715-730. 

MAKUSHOK, M. 1967. Whiptails (family Macrouridae or 
Coryphaenoididae Auct). Chapter IV. Pp. 200-227 in V. G. 
Kort (chief ed.), Biology of the Pacific Ocean, Book III, 
Fishes of the open waters. Moscow. (Transl. from Russian.) 
U.S. Naval Oceanogr. Off., Transl. 528, Wash., D.C. 

MARSHALL, NORMAN B. 1964. Bathypelagic macrourid 
fishes. Copeia 1964(1): 86-93. 

1965. Systematic and biological studies of the mac- 

rourid fishes Anacanthini-Teleostii. Deep-Sea Res. 12: 

299-322. 


nN 
in 
—_ 


1973. Family Macrouridae. Pp. 496-665 in Fishes of 
the western North Atlantic. Mem. Sears Found. Mar. Res. 
I(pt. 6). 

, AND A. VEDEL TANING. 1966. The bathypelagic 
macrourid fish Macrouroides inflaticeps Smith and Rad- 
cliffe. Dana Rep. 69: 1-6, pl. 1. 

MaulL, G. E., AND EINAR KOEFOED. 1950. On anew genus 
and species of macrourid fish, Phalacromacrurus panth- 
erinus. Ann. Mag. Nat. Hist., ser. 12, 3(35): 970-976. 

PARIN, N. V. 1975. Change of pelagic ichthyocoenoses along 
the section at the equator in the Pacific Ocean between 97 
and 155° W [in Russian, English summary]. Tr. Inst. 
Okeanol. 102: 313-334, figs. 1-6. 

, V. E. Becker, O. D. BORODULINA, AND V. M. 

TcHuvassov. 1973. Deep-sea pelagic fishes of the south- 

eastern Pacific Ocean [in Russian, English summary]. Tr. 

Inst. Okeanol., Akad. Nauk SSSR 94: 71-159. 

, G. N. POKHILSKAYA, Y. I. SAZONOV, AND B. I. 
FEDORYAKO. 1976. Rare and poorly known midwater 
fishes from the central and eastern equatorial Pacific Ocean 
{in Russian, English summary]. Tr. Inst. Okeanol., Akad. 
Nauk SSSR 104: 206-236. 

PEARCY, WILLIAM G., AND JULIE W. AMBLER. 1974. Food 
habits of deep-sea macrourid fishes off the Oregon coast. 
Deep-Sea Res. 21: 745-759. 

PECHENIK, L. N., AND F. M. TROYANOvVSKII. 1970. Trawling 
resources on the North-Atlantic continental slope. Main 
Administration of the Fishing Industry, Northern Basin, 
Murmansk, 65 pp. (Transl. from Russian.) Israel Progr. Sci. 
Transl., Jerusalem. 

Rass, T. S. 1963. Glubokovodnyye ryby—Dolgokhvosty 
(Pisces, Macruridae) Okhotskogo Morya. [Deep-sea gren- 
adier fishes (Pisces, Macruridae) of the Sea of Okhotsk. ] 
Tr. Inst. Okeanol., Akad. Nauk SSSR 62: 211-223. (Transl. 
from Russian.) U.S. Fish and Wildl. Serv., Seattle, Wash. 

Wyrrki, KLAus. 1967. Circulation and water masses in the 
eastern equatorial Pacific Ocean. Int. J. Oceanol. Limnol. 
1(2): 117-147. 


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PROCEEDINGS 


OF THE 


irs AY 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 8, pp. 253-265; 1 fig., 1 table. 


October 20, 1977 


LIZARDS OF THE GENUS LEPIDODACTYLUS (GEKKONIDAE) FROM 
THE INDO-AUSTRALIAN ARCHIPELAGO AND THE ISLANDS OF 
THE PACIFIC, WITH DESCRIPTIONS OF NEW SPECIES 


Walter C. Brown* 
Menlo College, Menlo Park, California 


and 
Fred Parker 


Wildlife Section, District Administration, Konedobu, Territory of New Guinea 


ABSTRACT: An annotated list and a key to species of the genus Lepidodactylus recorded from the Indo- 
Australian Archipelago are presented; four new species (L. magnus, L. mutahi, L. novaeguinea, and L. 
orientales) from New Guinea and the Solomon Islands are described. The relationships and distribution of the 


species are discussed. 


INTRODUCTION 


While revising Philippine species of the genus 
Lepidodactylus, one of us (Brown) examined 
samples, including types of extra-Philippine 
species. In the course of these comparisons, it 
became evident that the Indo-Australian species 
of Lepidodactylus, with the exception of the 
lugubris-woodfordi complex, comprise two 
groups distinct from the species in the Philip- 
pines. Also recent extensive collections from 
New Guinea and the Solomons include previ- 
ously undescribed species. It seems most ap- 
propriate to present these results in a report 
separate from our monograph, currently in prep- 
aration, on the gekkonid lizards of the Philip- 
pines. 


* Research Associate, Department of Herpetology, Cali- 
fornia Academy of Sciences. 


ACKNOWLEDGMENTS 


We are deeply indebted to Drs. A. G. C. 
Grandison of the British Museum (BMNH), Jean 
Guibe of the Muséum National d’Histoire Na- 
turelle, Paris (MNHN), K. Klemmer of the 
Senckenberg Museum (SM), W. R. Heyer of the 
United States National Museum of Natural His- 
tory (USNM), E. E. Williams of the Museum of 
Comparative Zoology (MCZ). H. Marx of the 
Field Museum of Natural History (FMNH), R. 
G. Zweifel of the American Museum of Natural 
History (AMNH), C. J. McCoy of the Carnegie 
Museum (CM), Paul Webber of the Australian 
Museum (AM), J. Menzies of the University of 
Papua New Guinea (UPNG), and N. Kwapena 
and M. C. Downes of the Department of Ag- 
riculture, Stock, and Fisheries, Papua New 
Guinea as curators for the Papua New Guinea 
Museum (DASF or PNGM), for the privilege of 


[253] 


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BROWN AND PARKER: INDO-AUSTRALIAN LEPIDODACTYLIDS 


examining pertinent material. CAS and CAS-SU 
refer to collections at the California Academy of 
Sciences. I am also deeply indebted to my col- 
leagues Dr. Alan E. Leviton and Mr. Robert C. 
Drewes for their assistance and suggestions. II- 
lustrations were prepared by L. Meszoly of the 
Museum of Comparative Zoology, Harvard 
University, and W. Zawojski of Stanford Uni- 
versity. This study is part of the Philippines- 
Pacific Islands research project under the aus- 
pices of National Science Foundation Grant 
GB-41947. 


SYSTEMATIC SECTION 


The available keys to Lepidodactylus , such as 
those of Taylor (1922) and de Rooi (1915) have 
not proven satisfactory to the non-specialists. 
For this reason, we have combined a more or 
less typical key to species-groups with diagnoses 
of species which should prove more useful. 


DIAGNOSTIC KEY TO SPECIES OF LEPIDODACTYLUS 
OCCURRING IN THE INDO-AUSTRALIAN 
ARCHIPELAGO AND ISLANDS OF THE PACIFIC 


(See Table 1 for additional data on species.) 


la. A relatively small species, adults usually 
less than 50 mm snout-vent length; ter- 
minal and 2-4 subterminal scansors on 
all digits but first divided (Fig. la); range 
of fourth toe scansors 11-15, covering 
distal 3/4 to 4/5 of toe; tail usually mod- 
erately flattened, with lateral fringe evi- 
dent (rarely subcylindrical); usually par- 
thenogenetic; males, when present in 
populations, with about 20-30 preanal 
and-temonal: pores’ ei ee 
L. woodfordi-lugubris complex 
1b. Terminal scansor on all digits entire or 
slightly notched (Fig. 1b); subterminal 
scansors entire or a few distal ones di- 
vided; tail subcylindrical without lateral 
(Rolie 2c es Sis io er oe eee 2 
2a. One or more (usually 2-4) subterminal 
scansors divided medially (Figs. 1b and 


il(cl)" ae tau Chee oie =e eee ere ee ee 3 
2b. All scansors entire, or at most, with only 

shallow median notches (Figs. Ic and 

1) cde Ey ee ee eae ee 6 


3a. Enlarged scales of pore-series and pores 
limited to preanal region or including 
only 1 or 2 scales on proximal part of 
thigh; no greatly enlarged scales on rest 


3b. 


4a. 


4b. 


SiGe 


of thigh; usually 20 or fewer preanal 
POFesvintmales <8 Sak 2 ee Pe 
Enlarged scales of pore-series not lim- 
ited to preanal region, at least one or 
more rows of scales on thigh distinctly 
enlarged although they may be sepa- 
rated from preanal series; 25 or more 
preanal and femoral pores in males_____- 
An intermediate-sized species, adults 
about 40-55 m snout-vent length; 16-19 
fourth toe scansors, covering toe to base 
(Fig. 1d); 11-13 first toe scansors; 4 to 
Ys webbed between third and fourth 
toes; 18-20 enlarged scales in preanal 
pore-series, bearing 16 pores for one 
male; brown blotches on head and nape 
(sharply outlined for Wild Island speci- 
mens), length of hind limb more than 
80% of axilla-groin distance __ L. pulcher 
A small species, adults about 36-45 mm 
snout-vent length; 11—16 fourth toe scan- 
sors, covering approximately distal %4 of 
toe; 8-11 first toe scansors; '/6 to '/4 
webbed between third and fourth toes; 
16 to 20 enlarged scales in preanal pore- 
series, bearing 12-19 pores for males; 
length of hind limb about 65 to 75% of 
axilla-groin distance; blotches on head 
and nape not sharply outlined 
L. novaeguineae new species 


. A relatively small species, 45+ snout- 


vent length (one available adult); 12 
fourth toe scansors, covering distal 34 
of toe; 11 first toe scansors; webbed 
only at base between third and fourth 
toes; the unique holotype, female, with 
about 38-39 enlarged preanal and fem- 
oral scales in position of pore-series, 
covering basal 34 plus of thigh 
BSS UE Ee Se teers L. gardeneri 


. A small to moderate-sized species, 


adults usually less than 50 mm in snout- 
vent length; 11-13 fourth toe scansors, 
covering distal 24 to 34 of toe (Fig. 1b); 
8-10 first toe scansors; about 4 to 3 
webbed between third and fourth toes; 
40-52 enlarged preanal and femoral 
scales in position of pore-series, con- 
tinuous and extending almost to distal 
end of thigh; bearing 39-52 pores (sam- 
plevor Ormales) 22 oe L. guppyi 
A relatively small species, about 36 mm 
snout-vent length for known adult speci- 


255 


256 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 8 
men; digits moderately dilated; 11+ snout-vent length (12 adults); digits rela- 
fourth toe scansors, 2 subterminal ones tively short and broadly dilated; 11-13 
divided; toes '/5 to '/4 webbed; unique entire fourth toe scansors, covering the 
holotype (a male) with about 34 enlarged distal 2 to % of the toe (Fig. le); 4 
preanal and femoral scales in the posi- to ¥2 webbed between third and fourth 
tion of the pore-series, continuous, toes; 12-15 enlarged scales in preanal 
beakine 52 pores: ere ae. L. shebae pore-series, separated by several scales 

6a. Adult size relatively small; enlarged (7 to 12 in samples) from moderately 
scales of pore-series and pores limited enlarged femoral series; bearing 11-14 
to the preanal region or with only 2 or preanal pores separated from femoral 
3 on proximal part of thigh; usually 20 or series of 10 to 14 in males; length of hind 
fewer preanal pores in males; webbed limb about 59 to 70% of axilla-groin dis- 
at base to '/s between 3rd and 4th toes 7 1 0 Ce <a aR rere J >: L. pumilus 

6b. Adult size variable; 26-50 or more en- 10b. A moderately large species, adults from 
larged preanal and femoral pore scales about 40 to 55 mm snout-vent length; 
in the position of the pore-series, in digits moderately dilated; 10 or 11 entire 
either a continuous series or separate fourth toe scansors, covering approxi- 
preanal and femoral series; 25 or more mately the distal 24 of toe; about 4 
preanal and femoral pores in a contin- webbed between third and fourth toes; 
uous or separate series in males__--____- 9 28-36 enlarged preanal and femoral pore 

7a. Small to moderate-sized species, adults scales, extending almost to distal end of 
usually less than 50 mm snout-vent thigh; bearing a continuous series of 27— 
length; digits relatively broadly dilated; 34 pores in males; length of hind limb 
9-12 undivided fourth toe scansors, usually more than 70% of axilla-groin 
covering distal 34 of toe; 8-10 first toe distance === L. mutahi new species 
scansors; webbed at base or to !/6 be- lla. A relatively large species, 53 to 57 mm 
tween third and fourth toes; usually 18 snout-vent length (3 adults); digits mod- 
or fewer scales in position of pore- erately dilated; 12-14 entire fourth toe 
series, limited to preanal region or with scansors; 9 or 10 first toe scansors; 
only one or two such scales on base of webbed at base only between third and 
thigh; bearing 15 or fewer preanal pores fourth toes; 32-36 enlarged preanal and 
in males; (see note under Discussion) __ 8 femoral pore scales in continuous series, 

7b. A relatively small species, 37 to 43 mm extending over proximal % of thigh; 
snout-vent length (3 adults); digits long bearing 28-30 pores in males______ L. oorti 
and relatively slender; 10-12 undivided 11b. A large species, adult males measure 50 
fourth toe scansors, covering approxi- to 71 mm snout-vent length; 12-15 entire 
mately distal *%4 of toe; 20-24 enlarged fourth toe scansors (Fig. 1c); 10-12 first 
scales in pore-series, confined to the toe scansors, '/s to '/; webbed between 
preanal region or only a few on base third and fourth toes; 40-50 enlarged 
of thigh; bearing 19 preanal pores (sam- preanal and femoral pore scales in a con- 
plevolvone male) 2.2... 2. sei tinuous series, extending almost to dis- 
ee a L. orientalis new species tal end of thigh; bearing 38 to 49 pores 

8a. Limited to Fiji Islands in central Pacific im males... =.= L. magnus new species 
Oceanke# Be. 26 vik Je ae L. manni 

8b. Limited to Christmas Island in Indian 
Ocean ee L. listeri LEPIDODACTYLUS SPECIES OF THE 

9a. Usually 4 or more webbed between INDO-AUSTRALIAN ARCHIPELAGO AND Is- 
third and fourth toes; usually 12 or fewer LANDS OF THE PACIFIC 
scansors beneath the fourth toe__________ 10 ; . 

9b. Usually only '/s or less webbed between Lepidodactylus gardener! Boutengcy 


Lepidodactylus gardeneri BOULENGER, 1897:306 (type- 


third and f : ‘ ; 
d ourth toes; usually more than locality: Rotuma Island, Polynesia; type in BMNH). 


12 scansors beneath the fourth toe______ 11 


10a. A relatively small species, 34 to 48 mm MATERIAL EXAMINED.—BMNH 1946.8.22.35 (holotype). 


257 


BROWN AND PARKER: INDO-AUSTRALIAN LEPIDODACTYLIDS 


Ficure 1. Inferior view of foot of five species of Lepidodactylus: (a) L. lugubris; (b) L. guppyi; (c) L. magnus; (d) L. pulcher; 


(e) L. pumilus. 


RANGE.—The known range is the type-local- 
ity, Rotuma Island. 


Lepidodactylus guppyi Boulenger 


Lepidodactylus guppyi BOULENGER, 1884:210 (type-locality: 
Faro Island, Solomon Islands; holotype in BMNH). 


MATERIAL EXAMINED.—BMNH _ 84.3.2.54 (holotype); 
1972.223-4, 1973.218; CAS 139650; MCZ 65862, 67122, 67124, 
67126, 74517-19, 75904 ab, 75905, 115559, and 115563-65. 


RANGE.—This species has been recorded 
from Bougainville, Buka, Faro, Guadalcanar, 
Kolambangara, Malaita, Puruata, Savo, Ster- 
ling, and Tulagi Islands in the Solomons. 
Hediger (1934:490) also lists L. guppyi as occur- 
ring in New Britain. 


Lepidodactylus listeri (Boulenger) 


Gecko listeri BOULENGER, 1888:535 (type-locality: Christmas 
Island, Indian Ocean; syntypes in BMNH). 
Lepidodactylus listeri KLUGE, 1967:9 


MATERIAL EXAMINED.—BMNH_ 1946.8.25.91 (holotype), 
1905.6.24.16-17, 1909.3.4.7; MCZ 143838; CAS 16861. 


RANGE.—This species is known only from 
Christmas Island in the Indian Ocean. 


Lepidodactylus lugubris (Dumeril and Bibron) 


Platydactylus lugubris DUMERIL AND BIBRON, 1836:304 
(type-locality: Otaiti (= Tahiti) Island, Polynesia; holotype 
in MNHN). 

Peropus neglectus GIRARD, 1858:197 (syn. fide M. A. Smith 
1935; type-locality: Rio de Janeiro, Brazil; holotype lost). 

Hemidactylus meijeri BLEEKER, 1859:47 (syn. fide M. A. 
Smith 1935; type-locality: Bintang, Rhio-Archipel fide M. A. 
Smith; holotype in BMNH). 

Gecko moestus PETERS, 1867:13 (syn. fide M. A. Smith 1935; 
type-locality: Palau Islands; holotype in Berlin Museum). 
Gymnodactylus caudeloti BAVAY, 1869:13 (syn. fide M. A. 
Smith 1935; type-locality: New Caledonia; repository data 

not available). 

Peripia mysorensis MEYER, 1874:129 (syn. fide M. A. Smith 
1935; type-locality: Mysore (= Biak Island); holotype in 
Berlin Museum). 

Peripia ornata MCLEAyY, 1878:98 (syn. fide Kluge 1963; type- 
locality: Port Moresby, New Guinea; type lost). 

Lepidodactylus divergens TAYLOR, 1918:242 (syn. fide M. A. 
Smith 1935; type-locality:Great Govenen Island, Sulu Is- 
lands, Philippines; type lost). 


MATERIAL EXAMINED.—MNHN _ 5323a-b_  (syntypes); 
AMNH 65484-87, 66223, 69628, 41867, 41875—-77, 41880-87; 
CAS 50274-96, 60613-18, 62485-86, 72031; CAS-SU 9159-60, 
9173, 9175-76, 9178, 9189, 1216466, 12176-81, 18047-56, 
one hundred plus uncatalogued; DASF 10297-99, 11462, 
11853-60; FMNH 44504, 73377 (paratype, L. divergens); MCZ 
16403, 19646, 27936, 33518, 48625-31, 49496-97, 69499, 
6950la-b, 26087-88 (paratypes, L. divergens), 85747-5S0, 
26091-93, 135431-32, 137648-50, 140981; USNM 28239, 
6804446, 68048, 68866, 70736, 119195, 121387, 121398. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 8 


RANGE.—This species has a wide distribution 
in the tropics, extending from India, Sri Lanka 
(Ceylon) and nearby islands through southeast 
Asia, Philippines, Indo-Australian Archipelago, 
northern Australia, islands of the tropical 
Pacific, and western Mexico and Central 
America. 


Lepidodactylus magnus, new species 


Holotype: MCZ 101504, an adult male, collected by J. Hig- 
ginbotham in the Kol Jim Valley Highlands, Papua New 
Guinea, 28 Mar. 1967. 

Paratypes: Central mountain area of New Guinea between 
about longitude 142° and 146° E, Lufa area: MCZ 96907-19; 
Karimui area: MCZ 101505—07, 101509; Nondiri area: MCZ 
91602-06, AMNH 98513-14; Yandime area: MCZ 91585-88, 
AMNH 98512; Orumba area: MCZ 91589; Yangamug]l area: 
MCZ 91597; Chauve area: MCZ 91598-99; Elmagale area: 
MCZ 91601, CAS 139656; Huon Peninsula: MCZ 54246; Kun- 
diawa area: 109242; Irumbafoie area: MCZ 91591-96, BMNH 
1974.4217, AMNH 95209-13; Kaironk Valley: UPNG 340S— 
17, CAS 139654-55, FMNH 197933, CM 59037, MNHN A833, 
USNM 198147; Gono: AMNH 92527-43 + 18 uncatalogued, 
CAS 139830; Kassam: AMNH 92653; Lalang: AMNH 95655; 
Rugli: AMNH 105880-92; Kwi Valley: MCZ 91597. 


DiaGnNosis.—A large Lepidodactylus , snout- 
vent length about 50 to 70 mm for adults; digits 
long, moderately dilated, Gekko-like; distal two 
thirds or more of ventral surface bearing undi- 
vided scansors, though one to four subterminal 
ones may occasionally exhibit shallow notches; 
12 to 13 scansors beneath fourth toe; 10 to 12 
beneath first toe; toes webbed at base between 
first and second toes and about one-fourth be- 
tween third and fourth toes; adult males with 
about 40 to 50 preanal and femoral pores, 
femoral series extending from base to near the 
distal end of thigh; sometimes disrupted within 
femoral series or between preanal and femoral 
series by one or a few scales; tail subcylindrical, 
without lateral flange or spines. 

DESCRIPTION.—A large Lepidodactylus, 
snout-vent length 50 to 70+ mm in 12 mature 
males (one 43-mm male immature), 55 to 67 mm 
in 10 mature females, and 24.5 to 25.9 mm (about 
25 to 27 mm before preservation) in four hatch- 
lings; moderately depressed, neither strongly 
robust nor slender in appearance; snout slightly 
tapered, rounded, length 38 to 41+% of length 
of head; internasal distance 46 to 49% of snout 
length; head breadth 72 to 80% of head length 
and 17 to 20% of snout-vent length; diameter of 
eye about 80 to 88% of snout length; rostral 
about 1-’% times as broad as long, somewhat 
longer between supranasals than at lateral mar- 


BROWN AND PARKER: INDO-AUSTRALIAN LEPIDODACTYLIDS 259 


gins; nostril surrounded by rostral, first upper 
labial, supranasal and two enlarged scales 
dorso-posteriorly; supranasals separated by two 
or three scales (most frequently three) bordering 
rostral; 9 to 12 upper labials, eighth to tenth be- 
neath median plane of eye; 9 to 10 lower labials, 
anterior ones as large as triangular mental; first 
row of scales behind mental gives appearance of 
four short chin shields followed by two to three 
rows of slightly enlarged scales; scales on snout 
larger than those on dorsal and lateral surfaces 
of body, which are covered by small, relatively 
uniform granular scales, enlarged tubercles lack- 
ing; limbs moderately developed, length of ex- 
tended hind limb (in preserved condition) 75 to 
84% of axilla-groin distance; toes webbed at 
base between first and second and as much as 4 
between third and fourth; digits moderately di- 
lated throughout their length (Gekko-like); 
breadth of fourth toe about 35 to 42% of its 
length to base of web; 10 to 13 (most frequently 
12) entire scansors under fourth toe, covering 
approximately distal % to 34 of toe (Fig. Ic); 10 
to 12 scansors under first toe; all digits but first 
finger and toe clawed; compressed, clawed 
phalanges free only at end of dilated part, ex- 
tending short distance beyond; a series of about 
50 enlarged scales in position of preanal and 
femoral pore-series, bearing 40 to 51 pores in 
mature males, preanal series forming a broad, 
rounded arch and femoral series extending al- 
most to distal end of thigh; (femoral series often 
exhibits some breaks of one to three or four 
scales without pores); pore-series followed by 
several rows of enlarged scales in preanal region 
but only by a single row on thighs before meet- 
ing smaller, more granular scales of posterior 
surface; tail appears subcylindrical, depth just 
posterior to basal swollen area 74 to 90% of its 
breadth at same point, and tail breadth 53 to 62% 
of head breadth; lateral margin without flange of 
skin or spines; scales on ventral surface of tail 
larger than on dorsal surface, squarish or at 
times fused to form quadrangular platelike 
scales (see also Table 1). 

Color. Jn preservative: dorsum and upper lat- 
eral surfaces grayish tan or brown, varying from 
relatively uniform or with a few dark lines or 
blotches along dorsolateral area to a pattern of 
five or six diffuse, wide, irregularly margined, 
transverse bands between nape and base of tail; 
venter rather uniformly light to heavily flecked 
with brown. Jn life: neck, body, and tail varying 


from pale grayish brown through dark gray- 
brown to brown; neck and body pattern often 
with shallow W-shaped bars; tail with pale 
blotches having distinct anterior border, fading 
posteriorly into darker ground color to next pale 
blotch; a pale stripe from ear to axilla and usu- 
ally from posterior corner of eye to ear; often a 
pair of dark stripes dorsolaterally on neck and 
usually a very distinct dark vertebral stripe on 
base of tail, reaching only to first or second pale 
blotch; frequently with pale spots around ear, 
and some specimens with pale spots on dorsum 
and in a line midlateral surface. 

Measurements (in mm) of holotype: snout- 
vent length 70.7; axilla-groin distance about 
34.5; hind limb length 27; length of head 16.7; 
breadth of head 13.8; snout length 6.7; diameter 
of eye 5.7; internasal distance 3.2; interorbital 
distance 3.3; tail breadth at base 7.3; tail depth at 
base 6.1. 

ECOLOGICAL Notes.—The localities listed for 
the holotype and paratypes are in the central 
mountains at elevations between about 1000 and 
2000 meters. One of us (Parker) and Professor J. 
I. Menzies have observed the species in crevices 
of limestone cliffs, among dead leaves, particu- 
larly of Pandanus and banana trees, and occa- 
sionally in houses, especially those built of bush 
material. These are resting sites during the day. 
The species is active at night. Pairs of eggs were 
found in a vertical fissure in a limestone cliff at 
Irumbafoie. Two large eggs were also present in 
the oviducts of several gravid females. 


Lepidodactylus manni Schmidt 


Lepidodactylus manni SCHMIDT, 1923:51  (type-locality: 
Suene, Viti Levu, Fiji Islands; holotype in MCZ). 


MATERIAL EXAMINED.—MCZ 16880 (holotype), AMNH 
81746-S1. 


RANGE.—This species is known only from the 
Fiji Islands. 


Lepidodactylus mutahi, new species 


Holotype: MCZ 127844, an adult male, collected by Fred 
Parker in Mutahi area, Bougainville Island, Solomon Islands, 
17 May 1966. 

Paratypes: Bougainville Island, same locality as holotype: 
MCZ 118317-118321; Tinputz area: MCZ 118316; Kieta area: 
MCZ 64152, 69216; Kunua area: MCZ 75863-75866, 75869, 
77590-77593, BMNH 1974.4218, UPNG 5192; Torokina area: 
USNM 120875-76, 120880, 120882, CAS 139657. 


DIAGNOosIs.—An intermediate-sized Lepido- 
dactylus, snout-vent length 37 to 56 mm for 


260 


adults; digits moderately dilated (Gekko-like); 
scansors covering approximately distal % of ven- 
tral surface of fourth toe; scansors undivided but 
often two to four subterminal ones with shal- 
low notches; toes webbed about '/s between the 
first and second and about 4+ between third and 
fourth; adult males with 29 to 34 preanal and 
femoral pores in a continuous series, extending 
almost to distal end of thigh; tail subcylindrical 
without lateral fringe of skin or spines. 
DESCRIPTION.—An intermediate-sized Lep- 
idodactylus, snout-vent length 37 to 56 mm 
for 16 mature males, 43 to 50 mm for five mature 
females; habitus moderately depressed, moder- 
ately slender; snout tapered, tip broadly 
rounded, its length about 39 to 42% of head 
length; internasal distance 40 to 50% of snout 
length; breadth of head 67 to 75% of head length 
and 17 to 20% of snout-vent length; diameter of 
eye 79 to 90% of snout length; rostral much 
broader than long and usually with a mid-dorsal 
bulge between the supranasals; nostril sur- 
rounded by rostral, first upper labial, suprana- 
sal, and two enlarged scales dorso-posteriorly; 
supranasals usually separated by two or three 
scales (rarely one) which border rostral; 10 to 12 
upper labials, usually ninth or tenth beneath cen- 
ter of the eye; 9 to 11 lower labials, anterior ones 
larger than triangular mental; latter followed by 
four, occasionally five, rows of enlarged scales 
in position of chin shields, anterior rows largest; 
scales on snout larger than those on dorsal and 
lateral surfaces of body which are covered by 
small, relatively uniform, granular scales; en- 
larged tubercles lacking; limbs moderately de- 
veloped, length of extended hind limb (in pre- 
served condition) 73 to 87% of axilla-groin 
distance; toes webbed, about '/5 between first 
and second, usually about 4 between third and 
fourth; digits moderately dilated, long; greatest 
breadth of fourth toe about 37 to 41% of its 
length to base of web; 9 to 12 undivided scansors 
beneath fourth toe, covering about distal % of 
toe; 11 to 14 under third toe and 8 to 10 under first 
toe (Table 1); all digits but first finger and toe 
clawed; compressed, clawed phalanges rising 
free at end of dilated part and extending for only 
short distance beyond; a series of 30 to 36 en- 
larged scales in position of pore-series; preanal 
series chevron-shaped, femoral series extending 
almost to distal end of thigh; a continuous series 
of 27 to 34 preanal and femoral pores in males; 
preanal series followed by several rows of en- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 8 


larged scales and preceded by two or three rows 
of slightly enlarged scales; femoral series form- 
ing an isolated enlarged row; tail subcylindrical, 
depth just posterior to basal swollen area 72 to 
92% of breadth at same point, and breadth 44 to 
51% of head breadth (one example greater than 
50%); lateral margins of tail without flange of 
skin or spines; scales on ventral surface of tail 
somewhat larger than those on dorsal surface 
and somewhat squarish. 

Measurements (in mm) of holotype: snout- 
vent length 55.2; axilla-groin distance 26.2; 
length of hind limb 19.8; head length 14.9; head 
breadth 9.8; snout length 5.6; diameter of orbit 
4.2; tail breadth 5.0; tail depth 3.6; internostril 
distance 2.2. 

Color. In preservative: dorsal ground color 
varies from gray-tan to brown, often with rather 
vague darker blemishes; venter uniformly light 
or with brown flecks which are most numerous 
laterally. Jn life: dorsum varies from pale 
yellow-brown to dark red-brown; often with a 
light stripe from eye to ear; usually orange 
patches on the tail; black blotches or bars which 
are evident on dorsum of many individuals ap- 
pear in life to vary for any given individual and 
are assumed to change in response to some 
stimulus. 

EcoLoGIcAL Note.—Field observations for 
the Kunua area (by Parker) indicate the species 
is most abundant in Pandanus in permanent 
coastal swamps, but that it also occurs in banana 
trees and newly cleared garden areas. 

RANGE.—Known at present only from Bou- 
gainville Island. 


Lepidodactylus novaeguineae, new species 


Lepidodactylus pulcher (part), LOVERIDGE, 1948:334; (part) 
WERMUTH, 1965. 


Holotype: CAS 89684, collected in Lake Sentani area, West 
Irian, by E. S. Herald, 1944. 

Paratypes: West Irian: BMNH 1938.6.6.72, FMNH 43059- 
60; Hollandia area, West Irian: AMNH 66346; Finchaven 
area, Marobe District, Papua New Guinea; CAS-SU 11028- 
29, 12182, USNM 11882425, 159826-27; Guisika area, Huon 
Peninsula, Papua New Guinea: USNM 119428, AMNH 6666S, 
66667, MCZ 49612; Mt. Nibo, Sepik District, Papua New 
Guinea: AMNH 100209-10. 


DiaAGNosis.—A moderate-sized Lepidodacty- 
lus distinguished by the following combination of 
characters: about 36 to 41 mm snout-vent length 
for 7 mature males; 11 to 14 scansors beneath 
the fourth toe, covering approximately the distal 


BROWN AND PARKER: INDO-AUSTRALIAN LEPIDODACTYLIDS 261 


24 of the toes; 8 to 11 beneath first toe (16 to 19 and 
10 to 13, respectively, for L. pulcher with which 
species has been confused); tail subcylindrical 
without fringe or spines; terminal scansor on all 
digits entire; a few subterminal ones, one to 
three on fourth toe divided; enlarged scales in 
pore-series limited to preanal region, about 14 to 
18 and bearing 12 to 18 pores in males; toes 
about 4 to 4% webbed. 

DESCRIPTION .—A relatively small Lepidodac- 
tylus, snout-vent length about 36 to 44 mm in 
adults; habitus not strongly depressed; snout 
slightly tapered and broadly rounded, its length 
about 35 to 39% of head length; internasal dis- 
tance 48 to 58% of snout length; breadth of head 
67 to 81% of head length, and 16.5 to 21% of 
snout-vent length; diameter of eye 85 to 97% of 
snout length; rostral about twice as broad as 
long, quadrangular to slightly trapezoidal; nos- 
tril surrounded by rostral, first labial, supranasal 
and two enlarged scales dorso-posteriorly; sup- 
ranasals usually separated by two scales which 
border the rostral; 9 to 12 upper labials, eighth or 
ninth beneath the center of the eye; 9 to 12 lower 
labials, the anterior ones larger than the triangu- 
lar mental; four or five rows of somewhat en- 
larged scales, in same position as chin shields, 
anterior row or two most enlarged; scales on 
snout larger than those on dorsal and lateral sur- 
faces of body, which are covered by relatively 
uniform granular scales; enlarged tubercles lack- 
ing; limbs moderately developed, length of ex- 
tended hind limb (in preserved condition) 66 to 
76% of axilla-groin distance; digits about !/¢ to '/3 
webbed, about '/s between first and second toes, 
and 4 to % between third and fourth toes; digits 
rather broadly dilated, noticeably more so in dis- 
tal half; breadth of fourth toe 37 to 40% of its 
length to the base of the web; terminal scansor 
on all digits entire; two or three subterminal 
scansors divided medially; 11 to 16 scansors be- 
neath fourth toe, covering about the distal %; 
all digits but first finger and toe clawed, com- 
pressed, clawed phalanges rising free at end of di- 
lated part and extending for only a short distance 
beyond; about 16 to 20 enlarged scales in posi- 
tion of pore-series, bearing 11 to 19 pores in ma- 
ture males (sample of three), limited to preanal 
region or with one or two at base of thigh; tail 
subcylindrical, depth just posterior to basal 
swollen area about 75 to 85% of breadth at a 
corresponding point, and tail breadth about 47 to 
58% of head breadth; lateral margin without 


fringes or spines; scales on ventral surface of tail 
larger than those on dorsal surface and more or 
less squarish in shape (see also Table 1). 

Color. In preservative: dorsal ground color 
rusty-tan to grayish, often with vague brownish 
markings; in some a little lighter in middorsal 
region and usually rather vague, reddish-brown 
flecks and markings on dorsal and dorso-lateral 
surfaces; or occasionally a woodfordi-like pat- 
tern of diffuse light and dark transverse bands; 
tail dorsally with a pattern of diffuse to fairly 
sharp, irregular, tan and reddish-brown cross 
bands. 

Measurements (in mm) of holotype: snout- 
vent length 40.1; axilla-groin distance 20.1; 
length of hind limb 15.4; head length 10.1; head 
breadth 8.1; snout length 3.9; diameter of eye 
3.3; tail breadth 4.5; tail depth 3.8; internasal 
distance 1.9. 

EcoLoGicAL Note.—Loveridge (1948:334) 
notes that one of the Gusiko specimens was col- 
lected on the trunk of a coconut palm and the 
other beneath the bark of a dead tree. 

RANGE.—Known from Hollandia area, West 
Irian to Huon Peninsula in northern Papua New 
Guinea. 


Lepidodactylus oorti (Kopstein) 


Gekko oorti KoOpsTEIN, 1926:77 (type-locality: Teun and 
Seruna Islands, Banda Sea and Samlakki, Tanimbar Is- 
lands; syntypes in BMNH, Leiden Museum and MCZ). 

Lepidodactylus oorti KLUGE, 1967:9. 


MATERIAL EXAMINED.—BMNH 1946.8.25.9, MCZ 38970 
and 39721 (3 syntypes). 


RANGE.—The species is known only from the 
Banda and Tanimbar Islands in the East Indies. 


Lepidodactylus orientalis, new species 


Gekko pumilus BOULENGER, 1898:697 (not Boulenger, 


1885a:473). 


Holotype: BMNH, 1897-12-10-7, collected in the Port 
Moresby area, Central District, Papua New Guinea, by Dr. L. 
Loria. 

Paratypes: Port Moresby area, Central District, Papua New 
Guinea: PNGM 10025, Konedobu, Central District, Papua 
New Guinea: MCZ 147357 and CAS 139833. 


The holotype of this previously undescribed 
species was one of three referred by Boulenger 
(1898) to G. pumilus. We have not seen the other 
two of Dr. Loria’s specimens, but we assume 
they are still in the museum at Genoa. 

DIAGNosis.—A relatively small Lepidodac- 
tylus distinguished by the following combination 


262 


of characters: snout-vent length 37 to 43 mm for 
four adults; digits moderately dilated (Gekko- 
like); 10 to 11 scansors covering % to nearly % 
of ventral surface of fourth toe; scansors undi- 
vided, but two or three subterminal ones often 
with shallow notches; about '/, webbed between 
third and fourth toes; adult males with a series of 
19 plus or minus preanal pores; enlarged scales 
in pore-series not extending beyond one or two 
scales on the base of the thigh; tail subcylindri- 
cal without lateral fringe of skin or spines. 
DESCRIPTION.—A relatively small Lepidodac- 
tylus, snout-vent length 37 to 43 mm in four 
adults; habitus moderately depressed, rather 
slender; snout tapered, tip broadly rounded, its 
length about 41 to 43% of head length; internasal 
distance about 50% of snout length; breadth of 
head 72 to 76% of head length and 18 to 19% 
snout-vent length; diameter of eye 69 to 86% of 
snout length; rostral broader than long, usually 
with a median, dorsal groove; nostril surrounded 
by rostral, first upper labial, supranasal, and two 
enlarged scales dorso-posteriorly; supranasals 
usually separated by two scales which border 
the rostral; 10 to 11 upper labials, usually the 
eighth or ninth beneath the center of eye; 9 or 10 
lower labials, anterior ones larger than mental; 
latter followed by three or four rows of enlarged 
scales in position of chin shields; scales on snout 
larger than those on the dorsal and lateral sur- 
faces of the body, which are covered by small, 
relatively uniform, granular scales; no enlarged 
tubercles; limbs moderately developed, length 
of extended hind limb (in preserved condition) 
67 to 78% of axilla-groin distance; toes weakly 
webbed, at base to about '/; between third and 
fourth toes; digits moderately dilated, long; 
greatest breadth of fourth toe about 40% of its 
length to the base of the web; 10 to 11 undivided 
scansors beneath the fourth toe; usually two or 
three subterminal ones slightly notched; scan- 
sors covering distal 3/5 to 7/3 of fourth toe; 8 or 9 
undivided scansors beneath the first toe (Table 
1); all digits but the first finger and toe clawed, 
the compressed, clawed phalanges rising free at 
the end of the dilated part, but extending only a 
short distance beyond; a series of about 20 to 24 
enlarged scales in position of pore-series includ- 
ing only one or two scales on basal end of thigh, 
and bearing, for males (one example), 19 preanal 
scales in a continuous series; the series followed 
by several rows of enlarged scales between it 
and vent; tail subcylindrical, depth just posterior 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 8 


to basal swellings 74 to 82% of breadth at same 
point, tail breadth 56 to 67% of head breadth; 
lateral margins of tail without flange of skin or 
spines; scales on ventral surface of tail larger 
than those on dorsal surface. 

Color. In preservative: dorsal ground color 
grayish tan to tan, with two or three pairs of 
small- to moderate-sized, brown, dorsolateral 
spots in region of fore limbs; dorsum, tail and 
limbs marked by faint to relatively prominent, 
narrow, irregularly margined, brownish, trans- 
verse bands, separated by somewhat broader 
light areas. 

Measurements (in mm) of holotype: snout- 
vent length 37.1; axilla-groin distance 19.5; 
length of hind limb 13.0; head length 9.1; head 
breadth 6.8; snout length 4.0; diameter of orbit 
3.2; tail breadth 3.8; tail depth 3.1; internasal 
distance 2.1. 

ECOLOGICAL NoTE.—We have no habitat 
data for this species. 

RANGE.—Known only from Konedubo and 
Port Moresby, Central District, Papua New 
Guinea. 


Lepidodactylus pulcher (Boulenger) 


Lepidodactylus pulcher BOULENGER, 1885b: 166 (type-locality: 
Wild Island, Admiralty Islands; syntypes BMNH). 


MATERIAL EXAMINED.—BMNH 1946.9.8.48-50 (syntypes); 
PNGM 10294, 10296, CAS 139832. 


RANGE.—This species is known from Wild Is- 
land off the northwest coast of Manua Island, 
Admiralty Islands, and Plot and Lengendrowa 
islands on the southeast coast. 


Lepidodactylus pumilus (Boulenger) 


Gecko pumilus BOULENGER, 1885a:473 (type-locality: Mur- 
ray Island, Torres Straits; type in BMNH). 
Lepidodactylus pumilus, KLUGE, 1967:9. 


MATERIAL EXAMINED.—BMNH _ 1946.8.25.93 (holotype); 
MCZ 135353-55, 13536466, 135529-30, 137568, 137586, 
139420, 14104849, 141366-68, AM 44233, PNGM 10293, CAS 
139834. 

RANGE.—This species is recorded from Mur- 
ray and Hammond islands, Torres Straits, and 
Daru Island of southeast New Guinea, Western 
District, Papua New Guinea. 


Lepidodactylus shebae (Brown and Tanner) 


Pseudogekko shebae BROWN AND TANNER, 1949:43 (type- 
locality: Lower Lunga River, Guadalcanar Island, Solomon 
Islands; type in the Museum, Brigham Young University). 

Lepidodactylus shebae, KLUGE, 1967:9. 


BROWN AND PARKER: INDO-AUSTRALIAN LEPIDODACTYLIDS 263 


MATERIAL EXAMINED.—Brigham Young University 7002 
(holotype). 


RANGE.—This species is known from Guadal- 
canar, Solomon Islands. 


Lepidodactylus woodfordi Boulenger 


Lepidodactylus woodfordi BOULENGER, 1887:334 (type- 
locality: Faro Island, Solomon Islands; type in BMNH). 
(See Discussion.) 


MATERIAL EXAMINED.—BMNH 1946.8.22.34. 


RANGE.—This questionable species has been 
recorded from the Solomon Islands, New 
Guinea, Philippines, and islands of the Pacific 
Basin. 


Lepidodactylus species 


Very small samples (1 to 3 specimens) from 
the following populations have not been referred 
to any of the designated species because of some 
differences in characters studied. We have not 
assigned these to a species pending the availabil- 
ity of larger samples. 

1. A unique specimen from Mount Riu on 
Sudest Island, Louisiade Archipelago, 
(AMNH 76766, female) is close to L. orien- 
talis in most characters but the interdigital 
webbing is more extensive. 

2. Two examples from the Adelbert Mountains 
in Madang District, Papua New Guinea, 
(AMNH 105087-88, male and female) appear 
most closely related to L. magnus based on 
the webbing and the shape of their digits but 
are most like L. pumilus in size and, possibly, 
in the pore pattern of males. 

3. Two specimens, BMNH_ 1974.3027 from 
Waigue Island and an uncatalogued example 
from Jappen Island, both off the northwest 
New Guinea coast, are close to L. 
novaeguineae in most characters but appear 
larger. 

4. Three specimens, MCZ 135433-34, 139418, 
mature males, from Ndrova Island, Admiralty 
Islands, are close to L. guppyi but are smaller 
(31.5 to about 38 mm snout-vent length) and 
have slightly fewer pores (34-36). 


DISCUSSION 


Kluge (1967) pointed out some of the 
similarities and differences among the related 
gekkonid genera Gekko, Pseudogekko, 
Lepidodactylus, and Hemiphyllodactylus. By 


numerical weighting (as a method of quantifying 
a judgment as to the degree of advancement ex- 
hibited for eleven characters analyzed), he 
ranked Pseudogekko, Lepidodactylus, and 
Hemiphyllodactylus in that order of change from 
an assumed Gekko-like ancestor. Kluge’s ap- 
proach enabled him better to define the genera 
and, in so doing, it became necessary to reassign 
several species within these genera, including 
some from Gekko to Lepidodactylus (Kluge 
1967:332). 

Russell (1972) included Gekko, Pseudogekko, 
Luperosaurus, Ptychozoon, Lepidodactylus, 
Hemiphyllodactylus, Gehyra, and Perochirus in 
a related group, the Gekko-group of the subfam- 
ily Gekkoninae. Within the genus Lepidodac- 
tylus, he regarded the evidence from the foot 
structure as supporting the hypothesis of an evo- 
lutionary trend involving a distal shift and reduc- 
tion in number of subdigital scansors, accom- 
panied by a median division of some of the distal 
ones to form two rows. We believe the evidence 
also indicates that a more depressed habitus 
parallels this shift as evidenced by the flattened 
and broadened tail and limbs. 

Based on these characters, we recognize three 
evolutionary lines within the genus. Species of 
Group I are characterized by numerous Gekko- 
like, undivided scansors on all digits. Members 
of Group II also have well developed scansors 
on almost all the undersurface of the digit, but a 
few subterminal scansors are divided. The tail in 
species of Group I and Group II is subcylindri- 
cal, without lateral flanges or spines. Species of 
Group III are characterized by a reduced 
number of scansors, with the terminal as well as 
a few subterminal scansors divided, and by 
being more depressed with flatter and broader 
tail. Three of the four new species of Lepidodac- 
tylus described here are members of the 
pumilus-oorti group (Group I) and one of the 
guppyi-pulcher group (Group II). Additional 
changes which are evident but more variable 
within these evolutionary lines involve breadth 
relative to length of dilated part of digit, the pro- 
portion of the toe covered by scansors, and the 
extent of webbing. 

We accept Russell's (1972:56) hypothesis that 
evolutionary trends within the genus involve 
shifts from the basic combination of characters 
exhibited by some Gekko-like ancestor. Group I, 
which is thus the most primitive of the three 
groups, includes seven species (Table 1). If we 


264 


assume that the Gekko-like ancestor was as large 
as nearly all living members of the genus Gekko, 
L. magnus would be the most primitive member 
in size, with L. oorti next. L. magnus also has 
long, slender digits and small webs between the 
digits. Based on these features, L. pumilus is the 
most advanced member of the group. The 
species is small and has extensive interdigital 
webbing, shorter limbs and digits, and more 
broadly dilated digits. The difference in adult 
size between L. magnus and L. pumilus is also 
evidenced by hatchlings. Four hatchlings of L. 
pumilus (MCZ 141048—49 and 1135529-30) were 
18.5-21.1 mm in snout-vent length, and four 
hatchlings of L. magnus (MCZ 98796-99) were 
24.5-25.9 mm in snout-vent length. 

The species of Group I occur in the islands of 
western Indonesia, New Guinea, and islands in 
the Torres Straits, the Solomons and Fijis in the 
Pacific, and Christmas Island in the Indian 
Ocean. L. pumilus of this group also is recorded 
from Hammond Island, offshore from Cape 
York, Australia, and may also occur in that re- 
gion of Australia. It is also of some interest that 
the two species within this group which are most 
alike are L. manni (Fiji Islands in the Pacific) and 
L. listeri (Christmas Island in the Indian Ocean). 
Based on the small samples available, the two 
are so similar in the characters studied that they 
cannot be distinguished except as widely sepa- 
rated populations (see Key). The two popula- 
tions may indeed be conspecific, but we have 
chosen not to synonymize them pending the 
availability of larger samples and field studies. It 
is also possible that recent human introduction 
accounts for one or the other of these popula- 
tions. 

Group II includes L. guppyi, novaeguineae, 
pulcher, shebae, and gardeneri—species with 
undivided terminal scansors on all digits, but 
with a varying number of subterminal ones di- 
vided. Within this group, L. gardeneri and L. 
guppyi comprise a closely related pair. The two 
differ most obviously in the extent of the devel- 
opment of interdigital webs (Table 1). L. pulcher 
and L. novaeguineae are another pair of related 
species which differ from each other in the 
number of scansors, the size of the eye relative 
to the snout length, and the length of the hind 
limbs relative to the axilla-groin distance (Table 
1). 

The species of Group II have a more restricted 
distribution. They are known from northern 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 8 


New Guinea, the Solomons, Admiralties, Bis- 
marks, and Rotuma Island north of the Fijis. 

Group III includes those species which are 
most advanced with respect to digital structure 
in terms of Russell’s hypothesis. Several distal 
scansors, including the terminal one, are di- 
vided, and scansors tend to be lost from the 
basal part of the digits and to be reduced in 
number. The tail is more depressed and is 
broader. 

The species of Group III include four endemic 
to the Philippine Archipelago and the widely dis- 
tributed L. lugubris-woodfordi complex. Popula- 
tions in this complex range from islands of the 
Indian Ocean and India to the western shores of 
the New World. The populations in Central 
America and South America, however, are ap- 
parently the result of very recent transport by 
man. In the past, some samples from various 
populations throughout the range have been re- 
ferred to L. lugubris by numerous authors, some 
have been described as distinct species only to 
be placed in synonymy by later authors, and still 
others from New Guinea, Philippines and other 
Pacific islands have been referred to L. wood- 
fordi. Boulenger’s (1887:334) brief description of 
L. woodfordi, based on a single specimen, was 
erroneously related to L. guppyi rather than L. 
lugubris. L. woodfordi and L. lugubris belong 
to the same evolutionary line and, in terms of 
characters used here, may possibly differ in 
color pattern and degree of flattening of the tail, 
but little else (Table 1). Adequate samples from 
numerous populations throughout the Pacific re- 
gion must be carefully studied and compared to 
determine the variability between and within the 
populations before the usefulness of these 
characters and the validity of these two named 
species, or the possible existence of additional 
races or species, can be finally determined. 

The distributional pattern of the four species 
which inhabit Papua New Guinea is also of in- 
terest because of the similarity in basic features 
with those seen in some of the Papuan species of 
the genus Sphenomorphus (Scincidae) such as 
solomonis, nigrolineatus, cinereus, and brun- 
neus (Greer 1973; Greer and Parker 1974). L. 
novaeguineae and §. solomonis are recorded 
only from low to moderate elevations along the 
north coast; L. magnus, S. brunneus, and S. 
cinereus are known from the central mountains; 
and L. orientalis and S. nigrolineatus are known 
from the Central District on the south coast, east 


BROWN AND PARKER: INDO-AUSTRALIAN LEPIDODACTYLIDS 265 


of the Gulf of Papua. The fourth species of 
Lepidodactylus, L. pumilus, is found in the 
Western District, west of the Gulf of Papua, and 
on the islands in the Torres Strait. 


LITERATURE CITED 


Bavay, A. 1869. Catalogue de reptiles de la Nouvelle 
Caledonia, et descriptions d’especes nouvelle. Mem. Soc. 
Linn. Normandie, Caen. 15: 1-37. 

BLEEKER, P. 1859. Reptilien en visschen van Bintang, aang- 
baden door E. Netscher, E. F. Meyer en H. Raet. Nat. 
Tijdschr. Neder.-Indié 16:45—47. 

BOULENGER, GEORGE A. 1884. Diagnoses of new reptiles 
and batrachians from the Solomon Islands, collected and 
presented to the British Museum by H. B. Guppy, Esq. 
Proc. Zool. Soc. Lond. 1884:210-213. 

1885a. Descriptions of three new species of geckos. 

Ann. Mag. Nat. Hist. Ser. 5, 16:473-475. 

1885b. Catalogue of lizards in the British Museum 

(Natural History). Vol. 1. Taylor & Francis, London. 

i-xii + 436 pp. 

1887. Second contribution to the herpetology of the 

Solomon Islands. Proc. Zool. Soc. Lond. 1887:333-338. 

1888. On the reptiles of Christmas Island. Proc. 

Zool. Soc. Lond. 1888:534-536. 

1897. On the reptiles of Rotuma Island, Polynesia. 

Ann. Mag. Nat. Hist., Ser. 6, 20:306-307. 

1898. An account of the reptiles and batrachians 
collected by Dr. L. Loria in British New Guinea. Ann. Mus. 
Civ. Stor. Nat. Genova, Ser. 2, 18:694-710, 3 pls. 

Brown, WALTER C., AND ANGEL C. ALCALA. 1964. Rela- 
tionship of the herpetofaunas of the non-dipterocarp com- 
munities to that of the dipterocarp forest of southern Negros 
Island, Philippines. Senckenb. Biol. 42:628-636. 

, AND Vasco M. TANNER. 1949. Rediscovery of the 
genus Pseudogekko with description of a new species from 
the Solomon Islands. Great Basin Nat. 9:41-45. 

DE Roots, NELLY. 1915. The reptiles of the Indo-Australian 
Archipelago. I. Lacertilia, Chelonia, Emydosauria. E. J. 
Brill, Leiden, i-xiv + 384 pp. 

DumerIL, A. M. C., AND GABRIEL BIBRON. 1836. Erpetologie 
général ou historie naturelle complete des reptiles. Vol. 3. 
Paris. i-iv + 517 pp. 

GIRARD, CHARLES. 1858. United States exploring expedition 
during the years 1838-1842 under the command of Charles 
Wilkes, United States Navy, 20 (Herpetology). i-xvii + 496 
pp., atlas 1-10 pp. + 32 pls. 

Gray, JOHN E. 1845. Specimens of lizards in the collection 


of the British Museum. Edward Newman, 
I-XXvVill + 289 pp. 

GREER, ALLEN E. 1973. Two new lygosomine skinks from 
New Guinea with comments on the loss of the external ear 
in lygosomines and observations on previously described 
species. Breviora 406: 1-25. 

, AND FRED PARKER. 1974. The fasciatus species 
group of Sphenomorphus (Lacertilia, Scincidae): notes on 
eight previously described species and descriptions of three 
new species. Proc. Papua New Guinea Sci. Soc. (1973) 
25:31-61. 

GUNTHER, ALBERT. 1864. The reptiles of British India. 
Taylor & Francis, London. i-xxvii + 452 pp. 

HeEpIGER, Henri. 1934. Beitrag zur Herpetologie und 
Zoogeographie Neu Britanniens. Zool. Jahrb. Abt. Syst. 
Oekol. Geogr. Tiere 65:389-582. 

KLUGE, ARNOLD G. 1967. Higher taxonomic categories of 
gekkonid lizards and their evolution. Bull. Am. Mus. Nat. 
Hist. 135:1-60. 

KopsTEIN, FELIX. 1926. Reptilen von den Molukken und der 
Benachbarten Inseln. Zool. Meded. Rijksmus. Nat. Hist. 
Leiden 1:71-112. 

LOVERIDGE, ARTHUR. 1948. New Guinean reptiles and am- 
phibians in the Museum of Comparative Zoology and 
United States National Museum. Bull. Mus. Comp. Zool. 
101:205—430. 

Meyer, A. B. 1874. Uber der von mir auf new-Guinea und 
den Inseln Jobi, Mysore und Mafoor im Jahre 1873 gesam- 
melten amphibien. Monatsber. Koenig. Preussischen Akad. 
Wiss. Berl. 1874:128-140. 

McLeEay, WILLIAM. 1878. The lizards of the ‘‘Chevert’’ Ex- 
pedition. Proc. Linn. Soc. N. S. W. (1877) 2:97-104. 

PETERS, WILHELM C. H. 1867. Herpetologische Notizen. 
Monatsber. Koenig. Preussischen Akad. Wiss. Berl. 
1867: 13-37. 

RUSSELL, ANTHONY P. 1972. The foot of gekkonid lizards: a 
study in comparative functional anatomy. Unpublished 
Ph.D. Thesis, University of London. i-iv + 376 pp. + 76 


London. 


pls. 
ScHMIpT, Kari. 1923. A list of Fijian lizards. Copeia 
1923:50-S2. 


SmitH, MALcotm A. 1935. Fauna of British India including 
Ceylon and Burma. Taylor & Francis, London. i—xii + 440 
pp. + 1 pl. 

TAYLor, EpwarpD H. 1918. Reptiles of Sulu Archipelago. 
Philipp. J. Sci. 13:233-267, 3 pls. 

1922. The lizards of the Philippine Islands. Philipp. 
Bur. Sci. Manila, Publ. 17: 1-269, 23 pls. 

WERMUTH, HEINZ. 1965. Liste der rezenten Amphibien und 
Reptilien: Gekkonidae, Pygopodidae, Xantusidae. Das 
Tierreich 80: 1-246. 


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PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 9, pp. 267-280; 3 figs., 5 tables. 


October 20, 1977 


ELEPHANT SEALS BREEDING ON THE 
MAINLAND IN CALIFORNIA 


By 


Burney J. Le Boeuf 


Crown College, University of California, Santa Cruz, California 95064 


and 


Kathy J. Panken 


Board of Studies in Biology and Coastal Marine Laboratory, 
University of California, Santa Cruz, California 95064 


ABsTRAct: A northern elephant seal gave birth on the mainland at Ano Nuevo Point, California, in 1975, the 
first mainland birth for this species in this century. Seven females gave birth in 1976 and 16 in 1977. The 
initiation of breeding in this location was apparently precipitated by crowding on the Ano Nuevo Island 
rookery, less than 1 km away. The new breeding area shows signs of increasing rapidly in size. Two hundred 
and forty-five thousand tourists attempted to view the seals during the last three years; 106,000 were taken on 
tours. The animals did not appear to be disturbed by the presence of humans in plain view at a distance of 10 


m or more. 


INTRODUCTION 


Northern elephant seals, Mirounga angus- 
tirostris, breed on offshore islands along the 
coast of California and Baja California, Mexico. 
Except when sick or injured, they have rarely 
been observed on the mainland during this cen- 
tury. However, the population has increased so 
dramatically during the last decade that space on 
islands in northern California has come to be in 
short supply. Male elephant seals began to ap- 
pear on the mainland directly across a narrow 
channel from the rookery on Ano Nuevo Island, 
California, during the 1967 breeding season 
(Poulter and Jennings 1966). They have been 
seen here in every subsequent breeding season 
and their number has steadily increased. In 
1975, afemale came ashore and gave birth on the 
mainland. To our knowledge, this was the first 


northern elephant seal born on the continental 
mainland during this century. The cyclic, pre- 
dictable appearance of these huge marine mam- 
mals on Ano Nuevo Point, a state reserve that is 
open to the public, has attracted thousands of 
tourists from nearby metropolitan areas. They 
have jammed the reserve in recent winters in an 
attempt to view and photograph the seals. Vari- 
ous confrontations between seals and people, 
ranging from humorous to dangerous, have oc- 
curred. 

The purpose of this paper is to trace the his- 
tory of elephant seals on the mainland in this 
area, to present data which enables us to specu- 
late on how and why breeding began here, and to 
discuss the implications of this extension in 
breeding range for these animals and man. 

To fully appreciate the recent appearance of 


[267] 


268 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 9 


elephant seals on the mainland, one must con- 
sider this development in the context of the 
species’ history over the last 175 years. 
Thousands of elephant seals existed at the be- 
ginning of the 19th century before sealing began. 
Their breeding range extended from Cabo San 
Lazaro in Baja California, Mexico, north to 
Point Reyes, California (Scammon 1874). How- 
ever, in the space of about 40 years, the elephant 
seal population was decimated by sealers who 
killed them for their oil. By 1860, the population 
was so severely reduced that sealing was no 
longer economically feasible and, in 1869, the 
species was considered virtually extinct. The 
nadir of the elephant seal population was around 
1894 to 1900 when less than 20 individuals sur- 
vived on a single beach on remote Isla de 
Guadalupe, an oceanic island approximately 300 
km off the coast of Mexico (Bartholomew and 
Hubbs 1960). 

In 1922, the species was given protection by 
the government of Mexico. The United States 
government gave similar protection a few years 
later. The number of elephant seals has been 
increasing logarithmically ever since. They now 
reoccupy most of their former range. Immi- 
grants from the parent colony on Isla de 
Guadalupe began to recolonize islands along the 
coast of Mexico and southern California during 
the period 1930 to 1950. Colonies in northern 
California were established on Ano Nuevo Is- 
land in 1961 (Radford, Orr and Hubbs 1965) and 
on Southeast Farallon Island in 1972 (Le Boeuf, 
Ainley and Lewis 1974). 

The Ano Nuevo Island rookery seems to have 
reached its carrying capacity. The number of 
pups born on Ano Nuevo Island has increased 
every year since the first two pups were born in 
1961 (Le Boeuf 1977). In 1977, 750 pups were 
born; at least this many females were present as 
well as 266 breeding-age males. During the peak 
of the breeding seasons in 1975, 1976 and 1977, 
the two sandy beaches on the island were 
crowded with females, their pups and adult 
males. Few additional males could have been 
accommodated on these preferred breeding 
areas (Le Boeuf and Briggs 1977). Evidently, 
some animals were affected by the crowded 
conditions and bred elsewhere. Many of the 
animals that colonized Southeast Farallon Is- 
land, 89 km to the north, during the period 1972 
to 1977, were immigrants from Ano Nuevo Is- 
land (Le Boeuf, Ainley and Lewis 1974; Harriet 


Huber, personal communication). Breeding 
began on the Ano Nuevo mainland during this 
same period. 


METHODS 


Ano Nuevo Island is located 30.6 km north of 
Santa Cruz, California. It is separated from the 
mainland point that juts out to sea by a 0.8-km 
channel. The study area is shown in Figure 1. 

Periodic censusing of the Ano Nuevo Main- 
land began during the 1968 breeding season (1 
December through 15 March) as an adjunct to 
systematic studies of the northern elephant seal 
on Ano Nuevo Island by Le Boeuf and col- 
laborators from the University of California at 
Santa Cruz (e.g., Le Boeuf and Peterson 1969; 
Le Boeuf and Briggs 1977). This paper covers 
censuses taken on the mainland during the 
period 1968 to 1977. 

The number of censuses varied greatly from 
year to year. The mean number of censuses per 
breeding season from 1968 to 1974 was 22 and 
the range was from seven in 1973 to 50 in 1971. 
Counts were made at various times of day, from 
late December until the first week in March, by 
walking the beach or from the island using a 
15-60 power spotting scope. In 1975 and 1976, 
censuses were taken one to three times a week, 
usually in the mornings between 0900 and 1100 
hours, by walking the length of the beach and 
searching the dune area. On several occasions, a 
second census was taken in the late afternoon. 
In 1977, censuses were taken daily at 0900 
throughout the breeding season. 

In our censuses, we distinguished males from 
females and adults from pups and juveniles. 
Breeding-age males were separated into four 
categories: adults (8+ years old), subadult or 
SA‘ (7-8 years old), SA* (6-7 years old), SA? (5 
years old) and SA! (4 years old). SA’ males are 
rarely observed on land during the breeding sea- 
son (Le Boeuf 1974). 

In addition to counting and categorizing the 
animals, we were able to keep a record of indi- 
viduals present. Studies on the island necessi- 
tated marking all males. This was done by 
bleaching a name or number into the pelage on 
both sides of each animal’s back as soon as it 
arrived (Le Boeuf and Peterson 1969) and by 
attaching numbered plastic tags into the interdig- 
ital webbing of the hindflippers (Le Boeuf, Ain- 
ley and Lewis 1974). Since most males moved 
back and forth between the island and the main- 


LE BOEUF AND PANKEN: ELEPHANT SEALS 


TUNITAS CREEK 


SAN GREGORIO CREEK 


PESCADERO CREEK 


BOLSA PT 


PIGEON PT 


FRANKLIN PT 


ARO NUEVO SOUTH BEACH 
POINT 


Afio Nuevo Island 
37°N.LAT. 06'05" 


Ano Nuevo 
Bay 


N 


aANO NUEVO IS © 


GAZOS CREEK 


PT_ANO NUEVO 
WADDELL CREEK 


SCOTT CREEK 


EL JARRO PT 


DAVENPORT 


gle xo 
ene 5 


s/f 


TO 
SANTA CRUZ 
30.6 km 


FiGureE 1. A schematic drawing of the Ano Nuevo mainland and Ano Nuevo Island. Stars indicate the location that females 
gave birth in 1976. Most of the females that gave birth in 1977 congregated in a harem on the east side of Ano Nuevo Point. The 


state reserve property is lined and the sand dunes are stippled. 


land, most seals observed on the mainland were 
already marked and could be identified individu- 
ally. When necessary, marking and tagging were 
also done on the mainland. In 1975, 1976 and 
1977, females and their pups were given indi- 
vidual marks (names) and tags denoting their re- 
lationship and the time and place of birth. Thus, 
in addition to determining the number and com- 
position of animals present each day, we could 
also identify individuals and follow them from 
day to day and from year to year. 

Since 1975, we have paid special attention to 
reproductive behavior and pup mortality. We 
recorded the following data on females: the time 


of arrival, the date and location of parturition, 
the sex and condition of pups, the frequency and 
date of each copulation as well as the male in- 
volved, and the date of departure. If an arriving 
female was tagged, its age could be determined. 
The location of parturition of all females was 
determined as well as their nursing behavior. We 
noted the reactions of females to their own pups 
and to those of other females. The incidence and 
manner of pup deaths were recorded. 


RESULTS 


Number. The number of elephant seals ob- 
served on the mainland during the breeding sea- 


270 


TABLE |. 
TEN-YEAR PERIOD. 


Breeding season 1969 1970 


Highest daily census 
of males on ML 7 6 3 10 


Number of different 
males on ML per 
season 7 11 W 15 


Number of different 
males on ANI per 
season 103 120 125 136 


Number of females 
on ML at peak 
season 0 0 0 0 


Number of females 
on ANI at peak 


season 178 219 291 296 


son increased dramatically from 1968 to 1977 
and coincided with a rapid increase in the size of 
the colony on the island. Table 1 shows that both 
the highest daily census and the number of dif- 
ferent males observed in a season increased sig- 
nificantly during the study period. No females 
were observed prior to 1975. The rapid growth of 
the Ano Nuevo Island population is reflected by 
the annual increase in number of females 
counted at peak season. The number of pups 
produced each year was approximately the same 
as the number of females counted at peak season 
(Le Boeuf and Briggs 1977). 

Distribution. Figure 1 shows the location on 
the mainland where the majority of seals were 
observed and the relationship of this area to the 
island. During the 1967 season, males were ob- 
served on Bight Beach and near Ano Nuevo 
Point (Poulter and Jennings 1966). In later sea- 
sons the seals occupied an increasingly larger 
area of the mainland with some individuals ap- 
pearing beyond the limits of the map shown in 
Figure 1. However, the majority of animals were 
observed between North Point and the eastern 
edge of South Beach; the heaviest concentration 
was usually in the immediate vicinity of Ano 
Nuevo Point. In recent years movements into 
the sand dune area became more frequent. On 
several occasions, males were observed as much 
as 0.5 km back into the dunes. In 1977, the 
dunes near Bight Beach were a gathering place 
for numerous subadult males. 

In 1972, males began hauling out on beaches 
beyond the boundaries of the state reserve. One 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 9 


NUMBER OF ELEPHANT SEALS ON ANO NUEVO MAINLAND (ML) AND ANO NUEVO ISLAND (ANI) DURING A 


1972 1973 1974 1975 1976 1977 
6 18 26 45 64 151 

16 38 44 94 148 320 
146 180 146 143 227 266 
0 0 0 1 7 16 
34] 360 426 556 687 740 


early excursion was recorded on national televi- 
sion; a mature bull surfaced on the Pebble Beach 
Country Club golf course, 72 km from Ano 
Nuevo Island, during the middle of a tourna- 
ment. He was back on the island the following 
day. In the same year, one subadult male made 
numerous trips to Southeast Farallon Island. He 
and two other males from the Ano Nuevo colony 
became permanent residents of this rookery the 
following year (Le Boeuf, Ainley and Lewis 
1974). In 1974, another bull took up residence 
for approximately three weeks on the main 
beach near the park entrance which borders the 
southern edge of the reserve. In 1975, 1976 and 
1977, males were sighted on several other 
beaches in the vicinity, extending from Bean 
Hollow Beach, 14.5 km to the north, to Daven- 
port Landing, 14.5 km to the south. Other public 
beaches where males were observed for several 
days include: Gazos Creek, Waddell Creek, 
Scott Creek, and Greyhound Rock (Figure 1). 
Most of the males observed on these peripheral 
beaches were mature, i.e., classified as an adult 
or near adult (SA‘). 

Prior to the arrival of females, spacing be- 
tween males on the beaches was 34 meters or 
more. Males were rather evenly distributed on 
Bight Beach, near Ano Nuevo Point, and on 
South Beach. When females began to give birth 
in the vicinity of Ano Nuevo Point (Figure 1), 
males began to concentrate in this area. This 
was especially noticeable in 1977 when a harem 
composed of 12 females formed in this area. 

Composition. Who were these males that fre- 


LE BOEUF AND PANKEN: ELEPHANT SEALS 


15 

th @ MATURE MALES 
13 

" X IMMATURE MALES 


FREQUENCY 


10-19 
Nov 9 Dec Dec Dec 


20-29 30 Nov- 20-29 30 Dec- 9-18 


8 Jan Jan 


19-28 29Jan 8-17 
Jan 7 Feb Feb 


TIME 


271 


18-27 28Feb- 9-18 19-28 
Feb 8March March March 


FiGuRE 2. The mean number of mature and immature males on the Ano Nuevo mainland calculated at 10-day intervals during 
the 1975 breeding season. A mature male is one who is approximately 8 years old or older. 


quented the mainland? Because nearly all of 
them were marked individually and daily rec- 
ords of their whereabouts were kept, it was evi- 
dent that males observed on the mainland were 
associated with the island population. Indeed, 
many males moved back and forth across the 
channel on a daily basis. For example, in 1976, 
65 percent of the males marked on the island 
were observed at least once on the mainland; all 
males observed on the mainland were also seen 
on the island. 

Table 2 shows that both mature and immature 
males were present on the mainland, with the 
former predominating in early years. This dis- 
crepancy may have been due in part to the fact 


that it was more difficult to mark all immature 
males because they continued to arrive through- 
out the season. In addition, a greater effort was 
made to mark adults in early years. On the is- 
land, the ratio of mature males to immatures was 
very close early in the study period. By 1975, 
immatures were more prevalent than matures on 
both the island and the mainland. The ratio of 
adults to immatures on the island in 1974 is in- 
consistent with other years because of a reduced 
effort to mark young males. In general, the age 
composition of males observed on the mainland 
reflected the age range of males that were pres- 
ent on the island. 

Mature males tended to outnumber immature 


TABLE 2. NUMBER OF MARKED MATURE (ADULTS AND SA‘ MALES, 7 YEARS OF AGE OR OLDER) AND IMMATURE 
BuLts (SA? AND SA® Mates, Less THAN 7 YEARS OLD) HAULED OuT ON ANO NUEVO MAINLAND AND ISLAND IN TEN 


CONSECUTIVE BREEDING SEASONS. 


Age 
Area Category 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 
Mainland Mature 4 9 5 10 23 31 46 55 58 
Immature 2. 2 2 6 aS) 11 48 93 262 
Island Mature 41 46 50 Sil 50 74 89 69 75 71 
Immature 48 56 64 60 77 92 49 74 152 195 


De PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 9 


50 


45 


40 @ MATURE MALES 


xX IMMATURE MALES 


FREQUENCY 


20-29 30 Nov.- 10-19 20-29 30 Dec- 


Nov 9Dec Dec Dec 


9-18 
8 Jan Jan 


19-28 19Jan- 8-17 18-27 28Feb- 9-18 19-28 
Jan 7 Feb Feb Feb 8 March March March 
TIME 


FIGURE 3. The mean number of mature and immature males on the Ano Nuevo mainland during 10-day intervals in the 1976 


breeding season. 


males during the beginning and end of each 
breeding season. The distribution of males on 
the mainland in 1975 and 1976 is shown in Fig- 
ures 2 and 3. This difference in distribution with 
age was due to multiple causes. One strategy of 
adult males was to rest on the mainland in De- 
cember and early January and then move to the 
island in late January when females began com- 
ing into estrus. Some individuals were very pre- 
dictable and repeated this pattern from one year 
to the next. Movement from the mainland to the 
island by the older, more dominant males was 
particularly evident in late January and early 
February. This pattern was less in evidence in 
1977. 

The peak number of immature males on the 
mainland occurred in mid-February and coin- 
cided with the time of highest copulatory activ- 
ity on the island (Figures 2 and 3). This pattern 
resulted in part from the fact that many young 
males were chased off the breeding beaches on 
the island by older, more aggressive bulls when 


the majority of females came into estrus. Thus, 
the young males were forced to haul out on vari- 
ous non-breeding areas on the island and on the 
mainland. This pattern of immature males reach- 
ing a peak on the mainland in mid-February was 
most obvious from 1974 to 1976. Figures 2 and 3 
also show that immature males arrived on the 
mainland later in the season, and they left earlier 
than adult males. 

Once females began to appear on the main- 
land, the most dominant males tended to remain 
near the females until the latter had given birth, 
nursed their pups, copulated and returned to 
sea. Only then did the males cross the channel to 
compete for females on the island. At season’s 
end, when all females had departed from the is- 
land, some males returned to the mainland to 
rest before going out to sea. 

Another strategy that was observed, particu- 
larly in early years, was for adult males to rest 
on the mainland for several days or even a few 
weeks after losing an important fight on the is- 


LE BOEUF AND PANKEN: ELEPHANT SEALS 273 
TABLE 3. REPRODUCTIVE SUCCESS OF MAINLAND-OBSERVED MALES ON ANO NUEVO ISLAND. 
Breeding season 1968 1969 1970 1971 1972 1973 1974 1975 1976 

Number of different 

males on ML per season 7 11 7 15 16 38 44 94 148 
Number of different 

ML-observed males 

that copulated on ANI D 4 4 4 6 12 14 39 34 
Percent copulations by 

ML-observed males on 

ANI 5 4 20 no data 19 31 6 68 42 


land. Several former harem masters on the is- 
land, deposed after commanding a harem con- 
taining hundreds of females for as long as six 
weeks, recuperated in exile on the mainland. 
The sight of bloody bulls prompted local resi- 
dents and tourists to refer to the mainland as 
‘“‘loser’s beach.’ This is misleading. Clearly, 
some males were resting after having been de- 
feated in a fight, but many of them on the main- 
land were by no means losers from the point of 
view of aggression or reproductive success. 
Table 3 shows that at least a quarter of the males 
observed on the mainland each year copulated 
on the island. In 1975 and 1976, males that had 
been seen on the mainland accounted for 42 and 
68 percent of the copulations observed on the 
island. Furthermore, in every year of the study, 
at least one male observed on the mainland was 
among the 10 highest ranking bulls on the island. 
The correlation between rank and reproductive 
success among males is high and positive (Le 
Boeuf 1974). Finally, we can reiterate that the 
majority of the males in the Ano Nuevo popula- 
tion were observed both on the island and on the 
mainland. 

As the total number of males on the mainland 
increased over the course of the study period, 
males began to arrive earlier and stay later in the 
season each year. In 1968, the first male arrived 
on 11 January and the last one left on the first of 
March. In 1977, the first male arrived on 15 
November and the last one departed on 26 
March. 

Non-reproductive behavior of males. There 
were no obvious differences in the behavior of 
mature and immature males on the mainland. 
Most of them spent their time sleeping on the 
dry sand near the upper edge of the beach. A few 
rested among the sand dunes. Threatening be- 
havior, chasing and an occasional fight were fre- 
quent activities of waking animals even before 


females began to appear. After the females ar- 
rived, aggressive encounters were most frequent 
in their vicinity. Young males were not as ag- 
gressive as older males and one might say that 
they ‘‘played”’ like adults in these aggressive 
confrontations. On several occasions, males at- 
tempting to land were rebuffed by the threats of 
dominant males on land—the result was that 
they landed further up or down the beach. This 
pattern of competition for social rank is typical 
of males on the island (Le Boeuf 1971, 1974). 
Many movements on the mainland consisted of 
simply going in and out of the water. Most often, 
this was in response to aggressive encounters 
with other males, but the animals also entered 
the water to cool off in hot weather. Males were 
often observed swimming along the surf line 
checking out the opposition, chasing off subor- 
dinates with the characteristic vocal threat, and 
remaining silent and low in the water when 
swimming past a dominant male. 

Duration of stay. The duration of each male’s 
stay on the mainland was variable. Some males 
were sighted on the mainland only briefly. 
Others crossed the channel several times a day 
and were sighted at several disparate locations. 
Some individuals remained in virtually the same 
location for several weeks. For example, in 
1974, an aging bull named Star caused us con- 
siderable trouble because he spent most of the 
breeding season sleeping on a narrow sandy ac- 
cess road to the beach, the route that we had to 
drive over in order to launch our boat to the 
island. The presence of females on the mainland 
after 1975 did not appear to change significantly 
the duration of the mainland haul-out for the 
majority of males; the exceptions were the few 
males who did most of the breeding. 

Reproductive activities on the mainland. On 
21 January 1975, a pregnant female hauled out 
near Ano Nuevo Point (Figure 1). Three days 


274 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 9 


later she gave birth to a female pup. A pregnant 
female had attempted to land here in 1973 but 
she was frightened off by tourists. The birth- 
place and age of the new mother were unknown. 
She and her pup were tagged and named, Bertha 
and Blue, respectively. Bertha nursed her pup 
daily for 30 days. A mature male, Baja, re- 
mained near here throughout the nursing period 
and kept all other suitors away. He alone, ap- 
parently, copulated with her on the same day 
that she returned to sea. All of these activities 
were similar to those seen on the island (Le 
Boeuf, Whiting and Gantt 1972), except that 
they were performed under the watchful eyes of 
thousands of tourists (e.g., San Francisco 
Chronicle, 29 January 1975; San Jose Mercury, 
18 March 1975). 

The 1976 breeding season. Bertha and six 
other females gave birth on the mainland in Jan- 
uary, 1976. Three additional females were ob- 
served and marked in the area early in the month 
but they remained for only a day or two before 
departing. 

The age and birthplace of only one mainland 
female was known. She was a three-year-old, 
born on Ano Nuevo Island and tagged (Green 
tag 1961) as a one-month-old weaned pup on 13 
February 1973. Her pup died when it was 11 
days old after being trampled by a charging bull. 
This female left the area later in the day. 

Besides the above, another female gave birth 
to a full-term stillborn. Thus, two pups died out 
of the seven born on the mainland. The other 
five pups survived to weaning and remained on 
the mainland for a few weeks after all adults had 
departed. 

An additional pup, a 1%2-week-old orphan 
found at Drake’s Bay near Point Reyes, was 
transported to the Ano Nuevo mainland on 26 
January by representatives of the California 
Marine Mammal Center, Fort Cronkhite, 
California. They placed the orphan alongside 
Bertha who was pregnant. She adopted the alien 
pup, allowing it to suckle along with her own 
pup who was born later in the evening. Both the 
adopted orphan and the filial pup survived to 
weaning age. The topic of adoption will be 
treated in more detail in a later section. 

In 1976, mainland females gave birth in three 
separate locations (Figure 1): three gave birth on 
the beach just north of Ano Nuevo Point, two on 
South Beach and two in the sand dunes, approx- 
imately 20 m inland from South Beach. Once 


females landed and gave birth they tended to 
remain in the same place throughout the 34 
week nursing period without ever entering the 
water. This is characteristic of all nursing 
elephant seals (Le Boeuf, Whiting and Gantt 
1972). The female whose pup was born dead was 
an exception. She was observed going in and out 
of the water several times as well as moving 
from one mainland harem to another. 

The most dominant male on the mainland was 
named Up and he spent most of the season near 
the females on Ano Nuevo Point. He alone 
copulated with both females that came into es- 
trus. The next two males in order of dominance 
were SRI-—236 and Atom. They positioned them- 
selves near the females on the South Beach and 
near those in the dunes, alternating positions 
from time to time, particularly early in the sea- 
son. Atom copulated with one female and SRI- 
236 copulated with two other females in the 
area. The three top-ranking males on the main- 
land were all adults. SRI-236 was ten years old, 
having been born and tagged on Ano Nuevo Is- 
land in 1966 (Poulter and Jennings 1966). He was 
next seen on the island during a breeding season 
in 1971. Up and Atom were approximately the 
same age, and they were first observed on Ano 
Nuevo Island in 1972. All three males returned 
to the island in subsequent breeding seasons and 
all of them were first observed on the mainland 
in 1974. The social behavior of other males on 
the mainland was concentrated around the 
breeding females and tended to be similar to that 
displayed by males competing on the island. 

The 1977 breeding season. The number and 
distribution of males and females on the main- 
land underwent an obvious change in 1977. The 
population more than doubled, and for the first 
time a harem formed on the mainland at Ano 
Nuevo Point consisting of 12 females at peak 
season. The number of males observed in the 
area was more than double that of the previous 
year, the increase being primarily in the subadult 
male category. The older, larger males concen- 
trated on Ano Nuevo Point near the female 
harem. Subadult males were more widely scat- 
tered except for a group of 50 to 60 individuals 
that congregated regularly in the sand dune area 
near the center of Bight Beach. 

Sixteen females gave birth on the mainland in 
1977, the original female, Bertha, being one of 
them. All births occurred during the interval 12 
January to 6 February. As in the previous year, 


LE BOEUF AND PANKEN: ELEPHANT SEALS 


275 


TABLE 4. INTERVALS IN Days BETWEEN FEMALE REPRODUCTIVE EVENTS ON THE ISLAND AND THE MAINLAND. 


Ano Nuevo Island 


Ano Nuevo Mainland 


1970! 19713 1976? 1976 1977* 
Arrival to parturition — 6.5 + 1.4 6.3 + 1.6 B251-= 1028 4.0 + 1.6 
(10) (29) (6) (14) 
Parturition to Ist 24.6 + 3.4 23.4 + 4.2 26.1 + 1.8 2338)-= 355 PULL SS a=: Ppp) 
copulation (9) (12) (25) (4) (13) 
Parturition to departure PAPAS) ae CNY) 29.0 + 5.0 77 fea) ae INS 26.0 + 2.7 PAS) se ila 
(nursing period) (12) (29) (48) (5) (14) 
Ist copulation to 3.31 2.8 S27 22 DY S\yil e= IE) 4.0 + 3.5 349 6s 2.3) 
departure (9) (11) (27) (4) (13) 
Arrival to departure 34.3 + 3.6 34.6 + 3.8 32.4 + 2.8 27.8 + 3.8 29° OEE 


(10) (9) 


(17) (4) (14) 


' From Le Boeuf, Whiting and Gantt (1972). 


? Unpublished data collected by B. Le Boeuf, K. Panken, M. Pierson and J. Reiter. 

* Excludes Green tag 3503 who gave birth and lost her pup. She did not nurse until 19 days later when she adopted a 
weaner. She departed 38 days after parturition and was on the mainland a total of 41 days. 

* Excludes two females who abandoned their pups, one on the day it was born and the other seven days later. They both 


went to sea on the day they abandoned their pups. 


several additional females landed early in Janu- 
ary but did not remain in the area long nor did 
they give birth. 

Besides Bertha, only one female from the pre- 
vious year returned to the mainland to give birth 
(Green tag 3503). Two females that gave birth on 
the mainland in 1976 gave birth on the island in 
1977 (Green tags 1861 and 3501). One female 
that gave birth on the mainland in 1977 (Green 
tag 1880) had given birth on the island in 1976. 
Of the three females that switched locations, 
two were three-year-olds who had lost their 
pups in the earlier year. 

Tags on four females revealed that they had 
been born on Ano Nuevo Island in 1973 and 
1974. Thus, three females were primiparous 
three-year-olds, and one of them was a four- 
year-old. 

Four females did not join the harem on Ano 
Nuevo Point (Figure 1). Bertha and an un- 
marked female gave birth in the dunes approxi- 
mately 100 meters east of the harem at Ano 
Nuevo Point. However, Bertha entered the 
harem site before leaving the rookery. A three- 
year-old female and another female (Green tag 
4543) gave birth on South Beach approximately 
50 meters east of the harem. 

There were no pup deaths on the mainland in 
1977. 

A male named BTLF (Green tag 3405) domi- 
nated all other males on the mainland, and he 
was involved in 60 percent of the matings ob- 


served. He was the first male to mate with 12 of 
the 16 females present. Most of the copulations 
by other males were performed while BTLF was 
engaged in combat with a challenger. BTLF was 
retagged on Ano Nuevo Island during the 1975 
breeding season; his earlier tag broke and could 
not be read. He copulated 29 times on the island 
in 1976 and was responsible for 4.7 percent of 
the matings observed. Up, the alpha male on the 
mainland in 1976, returned to the mainland in 
1977 but copulated only once. 

Female vital statistics. A comparison of inter- 
vals between various reproductive events on the 
mainland and the island reveals several differ- 
ences (Table 4). For nearly every comparison 
the interval is shorter for mainland females than 
for females giving birth on the island. Compared 
to island females, mainland females gave birth 
sooner after arriving, they copulated sooner 
after parturition, they nursed their pups at least 
one day less, and they spent less total time on 
land during the breeding season. All compari- 
sons between the 1976 island data and the 1977 
mainland data are statistically significant (t- 
tests, p < .05) except for the interval, first copu- 
lation to departure. 

Sex ratio of pups. Of the 24 pups born on the 
mainland during 1975 to 1977, 10 were males and 
14 were females. The sample size is too small to 
permit a meaningful test for a significant devia- 
tion from the 50:50 sex ratio. The sex ratio of 
pups born on the island has not deviated signifi- 


276 


TABLE 5. THE FREQUENCY OF VARIOUS TYPES OF 
NURSING RELATIONSHIPS OF FEMALES ON THE MAINLAND. 


Both 
years 
1976 1977 combined 
(N=7) (N= 16) (N=23) 
Nursed own pup to 
weaning age 
a. exclusively 2 6 8 
b. and an alien occasionally 0 3 3 
c. and an alien as well 2 5 7 
Nursed an alien after 
losing own pup 
a. exclusively (adoption) 1 0 1 
b. occasionally 
Did not nurse (because pup 
died, was abandoned, lost 
or stolen) D 2 4 


cantly from unity in recent years (Le Boeuf and 
Briggs 1977). It is notable that all four three- 
year-old females gave birth to female pups; a 
four-year-old female gave birth to a male. 

Maternal behavior. The most frequently ob- 
served nursing relationship in elephant seals is 
one in which the mother nurses her own pup to 
weaning and rejects the suckling attempts of 
alien pups. However, several deviations from 
this pattern occur (Le Boeuf, Whiting and Gantt 
1972). Indeed, complications in the nutritive re- 
lationship between females and pups on an 
elephant seal rookery were quite evident in the 
small and, hence, easily observed aggregation of 
females breeding on the mainland. Although it is 
beyond the scope of this paper to analyze this 
complex phenomenon in detail, we will describe 
some of these relationships to give the reader an 
appreciation for the range of possibilities ob- 
served. 

In 1976, an aggressive female with pup stole 
the newborn of a neighbor and nursed it, plus 
her own pup, to weaning age. The neighbor re- 
mained in the harem without nursing for 19 days 
and then adopted a weaned pup on the day its 
mother left the harem. She nursed the large 
weaned pup, a male, for 20 days. In contrast, 
two other females departed the area on the day 
their pups died. Bertha, as we have already men- 
tioned, adopted an orphan before she gave birth 
to her own pup; she nursed both pups to wean- 
ing age. Only two of seven females nursed their 
Own pups exclusively. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 9 


Additional complications were observed in 
1977. Although the majority of females nursed 
only their own pups, five females nursed an alien 
pup in addition to their own. One of them in- 
volved the serial adoption of two pups. While 
nursing her own pup, this female adopted an or- 
phan and nursed it for three days where upon 
she adopted another orphan, which she nursed 
for 12 days. Bertha, again, fostered a pup in addi- 
tion to her own. Two females that gave birth 
within one day of each other nursed each other’s 
pup as well as their own. Finally, two females 
abandoned their pups one and seven days after 
parturition, respectively. Their pups were nursed 
to weaning age by two different females with 
pups of their own. In both years, it was obvious 
to us that pups sharing a mother weighed less at 
weaning age than pups that suckled a female by 
themselves. However, weights were not ob- 
tained which would confirm this impression. 

The variety of nursing relationships observed 
on the mainland and the frequency of their oc- 
currence are summarized in Table 5. It is inter- 
esting to compare these figures with those col- 
lected on Ano Nuevo Island in 1970 and 1971 by 
Le Boeuf, Whiting and Gantt (1972). In this 
study, 72 percent of the females in a sample of 50 
nursed their own pups to weaning age (alone or 
in addition to an alien pup). This figure is similar 
to the figure obtained on the mainland in 1976 
and 1977, i.e., 78.3 percent of the 23 females 
nursed their own pups to weaning age. How- 
ever, the percentage of adoptions was more than 
twice as high on the mainland as on the island, 
34.8 percent of the females adopted a pup on the 
mainland compared to 14.0 percent on the is- 
land. We conclude that adoption occurs if a 
female nurses a pup consistently for one or more 
days regardless of whether she is already nurs- 
ing her own pup. The duration of fostering ob- 
served on the mainland ranged from 7 to 26 
days. Ten percent of the females in the island 
study did not nurse any pup after giving birth as 
compared to 17.4 percent in the mainland sam- 
ple. None of these females were ever observed 
copulating. Finally, in the larger population on 
the island, a few females each year permitted 
several pups to attempt to nurse at the same 
time. This permissiveness was not observed in 
mainland females, perhaps because there were 
fewer orphans present at all times. 

Non-breeding season observations. Prior to 


LE BOEUF AND PANKEN: ELEPHANT SEALS 


1976, the only observations of elephant seals on 
the mainland during the non-breeding season 
was in March and April. An occasional weaned 
pup swam across the channel and rested on the 
mainland for a few days, or a molting juvenile 
appeared for an equally short stay. However, in 
June 1976, many young males, who normally 
molt on the island at this time of year, were ob- 
served resting on the mainland at Bight Beach 
(Figure 1). For example, on 11 June, 20 males 
were resting on the mainland and 25 were in the 
water offshore. The highest number counted on- 
shore was 30 on 26 June (M. Bradeen, personal 
communication). Elephant seals are observed on 
Ano Nuevo Island at all times of the year (Le 
Boeuf, Ainley and Lewis 1974). Perhaps the 
presence of subadult males in June 1976 signals 
the beginning of a similar pattern on the main- 
land. 

The reaction of seals to humans. Prior to the 
1974 breeding season, the presence of elephant 
seals on the mainland attracted little attention. 
Only a few local residents and nature lovers 
knew about the seals and paid much attention to 
them. The distance of the seals’ resting place 
from the parking lot was also a deterrent. Ano 
Nuevo Point is about a 30-minute walk from the 
park entrance, and it can be cold, wet and windy 
in winter. However, just before the start of the 
1974 breeding season, a national magazine called 
attention to this unusual spectacle and recom- 
mended it to its readers (Sunset Magazine, De- 
cember 1973). Suddenly, this rather desolate 
area was flooded with hundreds of tourists. The 
understaffed and underfinanced Department of 
Parks and Recreation was not prepared to deal 
with this huge influx of people. During that win- 
ter, people were free to roam the beaches and 
dunes, and only the number entering the reserve 
was controlled. On one fair-weather weekend, 
as many as a thousand people strolled the 
beaches and sand dunes in the vicinity of Ano 
Nuevo Point. Each male elephant seal was sur- 
rounded by a circle of onlookers and amateur 
photographers. Families with small children 
walked too close to bulls weighing as much as 
2-3 tons, unaware of the danger. Some people 
gigged, prodded and threw pebbles at these 
giants in order to get more interesting photo- 
graphs. The public had not been informed that 
these animals might be dangerous or sensitive to 
human disturbance. In retrospect, it is surprising 
that no one was injured. We know of several 


277 


cases where people have been bitten by elephant 
seals at other places (the second author is one of 
them). 

It is equally surprising that the seals, all males 
that year, were so impervious to this constant 
attention and frequent harassment. Their typical 
response was to sleep and occasionally flip sand 
on their backs. When irritated, they gave a low- 
level open-mouth threat and increased the fre- 
quency of their sand flipping habit (Heath and 
Schusterman 1975). When pushed even further, 
some bulls reared up and bellowed the species- 
typical threat vocalization. Some charged the at- 
tacker for a distance of a few meters. Despite 
being provoked most males remained on the 
beach. Even those young males who backed 
down and were hounded into the water returned 
to the beach a short time later. Regardless of the 
throng of people in the area, the seals appeared 
to be primarily interested in each other’s move- 
ments. Several times it looked as if a male was 
charging a human when in reality he was attack- 
ing another male behind the person. When the 
seals were not being disturbed or approached 
too closely, they seemed to ignore the humans in 
the area. 

Disturbance to the seals in the park reserve 
was reduced considerably in subsequent years. 
Beginning in 1975, tourists were led on guided 
tours of the seals in groups of 20. Park rangers 
and student guides were informed about the 
animals, and were prepared to answer questions 
about them. A single path led through the sand 
dunes to overlooks near Ano Nuevo Point and 
South Beach where the incipient harems were 
located. Tourists were restricted to the paths 
and overlooks and were prohibited on the 
beaches and dunes where the animals were 
abundant. Approximately 245,000 people came 
to view the seals during the winter months of 
1975, 1976 and 1977; approximately 106,000 
people were led on tours. 

It is difficult to say whether a constant stream 
of tourists in plain view of the seals inhibited 
utilization of this area by the seals. Males did not 
appear to be affected very much even when 
harassed (a response that made them easy to 
slaughter in the last century). Females, on the 
other hand, are much more wary, especially 
when they first arrive and before they give birth. 
If disturbed during this period, females may 
reenter the water and either return to the same 
location later or give birth elsewhere. This pat- 


x PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 9 


tern has been observed on the island in previous 
vears (Le Boeuf, Whiting and Gantt 1972). We 
do not know whether females were frightened 
off by human activities on the mainland and, if 
so, the that were Since 
females often survey a potential breeding beach 
from the water before hauling out, this form of 
disturbance is difficult to monitor, Some females 
haul out and then leave. We know of at least 
three pregnant females that landed on the main- 
land in 1976 and subsequently departed before 
giving birth. A similar pattern was observed in 
1977. We do not know what caused these 
females to leave, and we were not able to deter- 
mine where they gave birth, 

Once a female gives birth, her movements de- 
crease significantly and she is not likely to aban- 
don her pup even if disturbed or approached 
closely by a human. Under these circumstances, 
a female will flip sand on her back nervously, 
issue a vocal threat and then attack. She will 
bite, hold and shake the victim. Female bites can 
produce severe injury. We saw several instances 
on the mainland where tourists ventured close 
enough to females and their pups to provoke 
arousal of females and the threat reaction just de- 
scribed. When advised, park rangers increased 
the viewing distance between seals and humans 
and this form of disturbance and potential 
danger was eliminated. When humans and seals 
were separated by approximately 10 m or more, 
the seals appeared to acclimate to the presence 
of humans in plain view, and females did not 
show obvious signs of alarm. As far as we could 
tell, the response of females to their pups was 
not affected adversely by human presence, but 
this subject deserves systematic study, 


numbers affected, 


DISCUSSION 


The initiation of breeding on the Ano Nuevo 
Mainland is a reflection of the continued growth 
and recovery of the northern elephant seal popu- 
lation following its near extinction in the last 
century. Indeed, the growth of the population 
has increased so rapidly of late that the Ano 
Nuevo Island breeding beaches have become 
filled to capacity at peak season (Le Boeuf and 
Briggs 1977). It is this condition which seems to 
have precipitated breeding in what might be 
called a suboptimal area—suboptimal because 
the mainland was formerly a habitat in which the 
seals were subjected to predation from carni- 


vores and coastal aborigines. Several lines of 


evidence lead to this conclusion: 1) females did 
not begin breeding on the mainland until the 
breeding beaches on the island began to be 
crowded (Le Boeuf and Briggs 1977); 2) during 
the same period, a new colony was formed on 
nearby Southeast Farallon Island, with most of 
the original settlers coming from Ano Nuevo Is- 
land; and 3) on Ano Nuevo Island, the first 
females to arrive at the start of the breeding sea- 
son have been observed preventing later arriving 
females from landing (Christenson and Le Boeuf 
1977) or forcing them to occupy peripheral posi- 
tions in harems where pup mortality is higher 
(Le Boeuf and Briggs 1977). 

Preliminary data suggest that it is the youngest 
females that begin breeding in new areas. Many 
of them are literally prevented from landing on 
breeding beaches or they are relegated to in- 
ferior locations by older, more aggressive 
females. Older females do this to enhance the 
probability of their own pups surviving (see Le 
Boeuf and Briggs 1977). A female is fully grown 
al six years of age; however, developing females 
usually give birth for the first time at age three, a 
few as early as age two. Subsequently, females 
give birth to a single pup annually (Le Boeuf, 
unpublished data). Most of the marked females 
breeding on Southeast Farallon Island from 1972 
to 1977 were between three and six years old. 
For example, four of six known-age females.that 
gave birth in 1975 were four years old, the other 
two were three years old and five years old, re- 
spectively. As late as 1977, seven females out of 
12 Known-age newcomers were three or four 
years old (H. Huber, personal communication). 
On the Ano Nuevo mainland, four of the 
known-age females that gave birth were three 
years old and one was four. Furthermore, ob- 
servations of known-age females on Ano Nuevo 
Island in 1976 by us, J. Reiter and M. Pierson 
revealed that over 50 percent of the females five 
years of age or less became separated from their 
pups, while less than 10 percent of the females 
SIX years of age or more lost their pups. Since 
mother-pup separation is the initial event in a 
syndrome culminating in pup mortality (Le 
Boeuf, Whiting and Gantt 1972; Le Boeuf and 
Briggs 1977), reproductive success, as measured 
by pup survival, is evidently lower in young 
females than in older females who are fully 
grown. Thus, it is obviously a reasonable strat- 
egy for young females to explore breeding in a 
peripheral place with more space, less competi- 


LE BOEUF AND PANKEN: ELEPHANT SEALS 


tion from older, more aggressive females and, in 
effect, a place where there is a higher probability 
that their pups will survive. 

Several discrepancies in behavior observed 
between females breeding on the island and 
those breeding on the mainland appear to be a 
consequence of the younger age of the latter. 
The shorter nursing period and reduced time 
spent on land during the breeding season by 
mainland females (Table 4) is consistent with the 
smaller size of young females. Small females are 
more severely limited by their body reserves 
while nursing (and at the same time, fasting 
completely) than larger females. Furthermore, 
the high percentage of adoptions we observed 
among mainland females seems to reflect their 
youth and inexperience. This relationship is sup- 
ported by observations of young females on the 
island. 

That only males were present indicates a 
wholly non-breeding use of the mainland prior to 
1975. The pattern of utilization by males was 
much like that which we observed in non- 
breeding areas on Ano Nuevo Island and in 
other rookeries; that is, it was a place for males 
to rest and a base from which they could make 
periodic forays into harems to enter into mating 
competition with other males. We consider the 
mainland as simply a new breeding area for the 
already established Ano Nuevo colony and 
stress that the mainland aggregation should not 
be considered a new colony. The establishment 
of a new colony develops in quite a different 
way. On Southeast Farallon Island, juveniles 
hauled out during the spring and fall for several 
years before adults began to appear in winter 
and breeding began. Females were the first to 
appear during the initial breeding season, just 
the opposite of the situation that obtained on the 
Ano Nuevo mainland (Le Boeuf, Ainley and 
Lewis 1974). 

A characteristic trait of most elephant seal 
females is to return to the same location to give 
birth year after year (Le Boeuf, Whiting and 
Gantt 1972). Because of this habit, the abun- 
dance of sandy beaches on the mainland, and the 
fact that the Ano Nuevo Island colony continues 
to increase in size despite the virtually complete 
utilization of breeding space on the island, we 
expect that the number of females breeding on 
the mainland will increase rapidly in the near 
future. During the last ten years, the mean 
annual increase in pups born on the island 


279 


was 16.9 percent with a range of 3 to 31 per- 
cent (Le Boeuf and Briggs 1977). Future in- 
crements in pup production within the colony 
will probably be borne by the mainland, es- 
pecially since the Southeast Farallon Island 
colony, which has received immigrants from 
Ano Nuevo Island for the past six years (Le 
Boeuf, Ainley and Lewis 1974; H. Huber, 
personal communication), is fast approaching 
its carrying capacity. In short, we expect that 
the size of the breeding unit on the mainland 
will increase very rapidly. If conditions are not 
altered drastically, we project that over 1,000 
pups will be born there within the next decade. 
The possibility of such a great increase in the 
number of seals breeding at this location will 
provide the public with a resource that is recre- 
ational, informative and unique. The abundance 
of tourists that have come to view these animals 
here in the past few years attests to public inter- 
est. Numerous busloads of children from 
elementary and high schools for miles around 
have taken field trips to this area. There is no 
other place in the United States where breeding 
pinnipeds can be viewed so easily and from such 
close quarters. State and federal officials will 
have to anticipate continued growth of this seal 
colony and prepare both for the protection of 
these animals and for full and imaginative utili- 
zation of this natural phenomenon as a resource 
of great interest and education for the public. 


ACKNOWLEDGMENTS 


We thank A. Hoover, J. Perlsweig, M. Pier- 
son, J. Reiter, M. Reidman, B. Tyler, several 
Afio Nuevo State Reserve rangers, and numer- 
ous student researchers and interns for field 
assistance; J. Anderson, Environmental Studies 
Internship Coordinator, University of California, 
Santa Cruz, for facilitating various aspects of 
data collection; and R. Werts, Manager, San 
Mateo Coast Area, Department of Parks and 
recreation, for permission to work in the reserve 
and for cooperation in all aspects of the study. 
This research was supported in part by National 
Science Foundation grant BNS 74-01363 A02 to 
B. Le Boeuf and authorized by Marine Mam- 
mal Permit No. 60, U.S. National Marine Fish- 
erles Service. 

LITERATURE CITED 
BARTHOLOMEW, G. A. AND C. L. Hugss. 1960. Population 


growth and seasonal movements of the northern elephant 
seal, Mirounga angustirostris. Mammalia 24:313-332. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 9 


CHRISTENSON, R. E. AND B. J. LE BoeEuF. 1977. Aggressive 
behavior of the female northern elephant seal. Behaviour (in 
press). 

HEATH, M. E. AND R. J. SCHUSTERMAN. 1975. ‘‘Displace- 
ment”’ sand flipping in the northern elephant seal, Mirounga 
angustirostris. Behav. Biol. 14:379-385. 

Le Boeur, B. J. 1971. The aggression of the breeding bulls. 
Nat. Hist. 80:82-94. 

1974. Male-male competition and reproductive suc- 

cess in elephant seals. Am. Zool. 14:163-176. 

. 1977. Back from extation? Pac. Discovery (in press). 

, D. G. AINLEY AND T. J. Lewis. 1974. Elephant 

seals on the Farallones: population structure of an incipient 

breeding colony. J. Mammal. 55:370-385. 

, AND K. T. BricGs. 1977. The cost of living in a seal 

harem. Mammalia (in press). 


, AND R. S. PETERSON. 1969. Social status and mating 

activity in elephant seals. Science 163:91-93. 

, R. J. WHITING AND R. F. Gantr. 1972. Perinatal 
behavior of northern elephant seal females and their young. 
Behaviour 43:121—156. 

PouLTER, T. C. AND R. JENNINGS. 1966. Annual report to 
the Division of Beaches and Parks, State of California. Stan- 
ford Research Institute, Menlo Park, California. 27 pp. 

RADFORD, K. W., R. T. ORR AND C. L. Husss. 1965. Rees- 
tablishment of the northern elephant seal (Mirounga angus- 
tirostris) off central California. Proc. Calif. Acad. Sci., ser. 
4, 31:601-612. 

ScAMMoNn, C. M. 1874. The marine mammals of the north- 
western coast of North America. John H. Carmany & Co., 
San Francisco. 


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PROCEEDINGS 
OF THE LIBRARI 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 10, pp. 281-295; 16 figs. 


October 20, 1977 


DESCRIPTIONS OF NEW SPECIES OF PERDITA (HYMENOPTERA, 
APOIDEA) IN THE COLLECTION OF THE 
CALIFORNIA ACADEMY OF SCIENCES 


P. H. Timberlake* 


Department of Entomology, Division of Biological Control, 
University of California, Riverside, California 92502 


ABSTRACT: Described from the western Nearctic are 16 new species of Perdita bees that belong to 4 subgenera 
and 3 groups as follows: in the subgenera Cockerellia (autumnalis and pachygnatha), Hexaperdita (cinctiven- 
tris, infuscata, and zavortinki), Perditella (pusillissima), and Perdita (Octomaculata group—esmeraldensis , 
inyoensis , medialis , pauliana, and willcoxiana; Ventralis group—diminutiva and gracilior; and Sphaeralceae 


group—apicalis , bifasciata, and perlucens). 


INTRODUCTION 


The genus Perdita consists of mostly small, 
colorful bees distinguished from other genera of 
panurgine bees by the very short, truncate mar- 
ginal cell and the short last submarginal cell in the 
forewings. The genus is noteworthy because of 
the great number of included species, several 
hundred being known, its relatively limited geo- 
graphical distribution, most of the species being 
restricted to the arid western United States and 
Mexico, and the oligolecty shown by most 
species. The genus was monographed in a series 
of seven parts that appeared in the University of 
California Publications in Entomology (Timber- 
lake 1954-1968). 

This report is based on a collection of Perdita 
received from the Academy in the spring of 1975 
and represents the accumulation of material over 
a period of some forty years. Four of the 16 new 
species are from New Mexico and the southeast- 


* Research Associate, Department of Entomology, Cali- 
fornia Academy of Sciences. 


ern part of Arizona, two from Esmeralda County, 
Nevada, and the remainder from California and 
Baja California. 


SPECIES DESCRIPTIONS 


Genus Perdita Smith 
Subgenus Cockerellia Ashmead 
Perdita autumnalis, new species 


This species of Cockerellia runs in the key 
(Timberlake 1968: 19) to luculenta Timberlake, 
but differs in the large dark mark on the clypeus, 
more strongly punctured frons and in having the 
abdomen black with three white bands. 

FEMALE.—Head and thorax dark green, pro- 
podeum nearly black. Mandibles, with exception 
of testaceous red apex, creamy white; labrum, 
clypeus, oblique lateral marks intruding between 
foveae and eye, transverse supraclypeal and 
rather small subantennal marks creamy white. 
Clypeus with large, triangular, median, black 
mark, narrowed above, and usually enclosing 
light spots. Line on collar, interrupted band on 


(281) 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 10 


hind margin of pronotum and tubercles white; 
thorax otherwise dark. Abdomen black, with 
small spot on lateral margins of tergite 1 and 
rather narrow band on tergites 2 to 4 white. Band 
on tergite 2 touching lateral foveae, that on tergite 
3 curved slightly backward at outer ends and 
reaching lateral margins, and that on tergite 4 
enclosed. Pygidial plate testaceous. Legs black, 
apices of front and middle femora, anterior sides 
and bases of tibiae broadly white, and tarsal seg- 
ments testaceous. Antennae dark, scape nar- 
rowly white beneath, flagellum more yellowish, 
but dusky above. Tegulae white at base and sub- 
hyaline on outer margin. Wings milky hyaline, 
nervures pallid, margins of stigma yellowish, 
subcosta fuscous. 

Head distinctly broader than long, cheeks 
strongly receding and three-fourths as wide as 
eyes, but narrowed anteriorly. Lateral ocelli 
about their distance apart from margin of eyes 
and occipital margin. Facial foveae strongly im- 
pressed, close to margin of eyes, nearly as wide 
as diameter of ocelli and reaching about three- 
fourths of distance from level of antennal sockets 
to level of anterior ocellus. Antennae inserted 
below middle of face, slightly clavate with middle 
joints of flagellum about as long as thick. Anten- 
nal sockets much closer to each other than to 
margin of eyes. Face below level of antennae 
convex, with clypeus prominent, disk of clypeus 
as broad as high and truncate at summit. Mandi- 
bles broadly dilated on inner margin, with in- 
curved acute apex. Proboscis much longer than 
head and much exceeding fossa in repose. Ptero- 
stigma as long, and about three-fourths as wide, 
as first submarginal cell; marginal cell longer be- 
yond than beneath stigma, with metacarpus half 
longer again than apical truncation. 

Head and thorax polished, frons with dense, 
strong, fine punctures, extending short distance 
below level of antennae, but clypeus with very 
few punctures. Punctures of mesonotum similar 
to those of frons but less dense, and those of 
mesopleura finer. Abdomen moderately shining, 
with dense, shallow punctures. Pubescence 
white, erect and rather dense on cheeks, sides 
and undersurface of thorax; somewhat shorter on 
mesonotum and still shorter on face above level 
of antennae. Apical fimbria of abdomen long and 
rather dense. Scopal hair of hind legs long, fine 
and very dense. 

Length, 6.5-7 mm; anterior wing, 5.5 mm; 
width of abdomen, 2.5 mm. 


HOLOTYPE FEMALE.—2.4 km (1.5 miles) SW of Wildrose 
Station, Death Valley National Monument, Inyo County, 
California, on Chrysothamnus paniculatus , 6 Nov. 1968 (P. H. 
Arnaud, Jr.). 

PARATYPES.—Three females, same data as type, and 12 
females, Water Canyon, Panamint Valley, Inyo County, 
California, 701 m (2300 ft), on same flower, 7 Nov. 1968 (P. H. 
Arnaud, Jr.). 

Types in collection of the California Academy of Sciences, 
San Francisco, and two paratypes in collection of the Univer- 
sity of California, Riverside. 


Perdita pachygnatha, new species 


In the key to Cockerellia (Timberlake 1954: 
386), the female falls in the group of species sepa- 
rated in couplets 21 to 24, agreeing best with 
‘‘albipennis ,"’ now called lingualis Cockerell, in 
sculpture and markings but differing from other 
species of Cockerellia in the short, broad mandi- 
bles with usual incurved apical part very short. 

FEMALE.—Head and thorax dark green, with 
propodeum more bluish. Mandibles, with excep- 
tion of reddish tips, black; labrum, broad median 
part of clypeus and supraclypeal area black; lat- 
eral third of clypeus, transverse lateral marks 
(somewhat widened on orbits), tubercles, and 
narrow, interrupted band on hind margin of pro- 
notum, creamy white. Abdomen black, with 
broad white band on tergites 2 to 4, curved back- 
ward at outer ends and reaching lateral margins 
on 2 and 3, shorter and straighter on 4, narrowly 
interrupted on 3 and 4, and moderately widely 
interrupted on tergite 2. Pygidial plate ferrugi- 
nous. Legs black with small joints of tarsi some- 
what reddish. Antennae black, flagellum broadly 
dark reddish ferruginous beneath. Tegulae tes- 
taceous, with white spot at base. Wings whitish 
subhyaline, nervures and stigma testaceous. 

Head as broad as long, with cheeks strongly 
receding; nearly as broad as eyes; lateral ocelli 
about their distance apart from margin of eyes 
and somewhat closer to occipital margin. Facial 
foveae well impressed, very close to margin of 
eyes and reaching about two thirds of way from 
level of antennal sockets to level of anterior ocel- 
li. Antennae inserted just below middle of face, 
with flagellum moderately clavate, with middle 
joints about as long as thick. Face below level of 
antennae strongly convex from side to side, with 
clypeus very prominent. Mandibles robust, 
hardly reaching beyond median line of head, with 
usual incurved apical part very short. Proboscis 
short, retracted in holotype, with apex of galeae 
reaching base of stipites. Pterostigma about half 


TIMBERLAKE: NEW PERDITA 


as wide as first submarginal cell; marginal cell 
somewhat longer beyond than beneath stigma, 
with metacarpus more than twice as long as api- 
cal truncation. Pygidial plate about as long as 
wide at base, with side converging to rather nar- 
row, rounded apex. 

Head and thorax shining, finely punctured. 
Frons very minutely tessellate, with fine 
punctures, about a puncture width apart, but 
leaving impunctate space in front of ocelli. 
Punctures of clypeus somewhat coarser and well 
separated, and those of mesoscutum also coarser 
and moderately dense. Mesopleura minutely tes- 
sellate, with fine separated punctures. Pubes- 
cence white, moderately long and dense on 
cheeks and thorax, and thinner on face. Apical 
fimbria of abdomen long but thin. Hair of legs 
white and abundant, with scopal hair of hind legs 
very long, fine and dense. 

Length, 6 mm; anterior wing, 4.5 mm; width of 
abdomen, 2.2 mm. 

HoLotyPe FEMALE.—2.4 km (1.5 miles) SW of Wildrose 
Station, Death Valley National Monument, Inyo County, 
California, 953 m (3100 ft), at flower of Chrysothamnus 
paniculatus, 6 Nov. 1968 (P. H. Arnaud, Jr.). 


Type in collection of the California Academy of Sciences, 
San Francisco. 


Subgenus Hexaperdita Timberlake 
Perdita cinctiventris, new species 


This species of Hexaperdita would fall with the 
species separated in couplets 22 to 25 in the key 
(Timberlake 1956: 249) and can be distinguished 
from foveata Timberlake, graenicheri Timber- 
lake, bishoppi  Cockerell, and _ boltoniae 
(Robertson) by having the face entirely dark and 
the light bands of the abdomen entire. 

FEMALE.—Head and thorax dark green with- 
out markings, except tubercles and anterior bor- 
der of pronotum white. Abdomen black, with 
broad, entire, yellowish-white band at base of 
tergites 2 to 5, that on tergite 2 touching lateral 
foveae, others not quite reaching lateral margins; 
tergite 1 with two small white spots, close to- 
gether at summit of basal declivity. Legs entirely 
dark. Antennae dark, flagellum brownish be- 
neath. Tegulae testaceous. Wings dusky hyaline, 
nervures dark, disk of stigma yellowish. 

Head moderately broader than long; cheeks 
receding, about three-fourths as wide as eyes. 
Lateral ocelli slightly farther apart than their dis- 
tance from margin of eyes. Facial foveae well 
impressed, and close to margin of eyes, about 


283 


four times longer than wide, and reaching about 
three fourths of distance from level of antennal 
sockets to level of anterior ocellus. Antennae 
inserted below middle of face, with flagellum cla- 
vate and middle joints broader than long. Face 
depressed, with clypeus gently convex, its disk 
broader than long and rounded at summit. Man- 
dibles reaching for margin of labrum, parallel 
sided and evenly rounded at apex. Proboscis 
moderately long, galeae shorter than stipites. 
Pterostigma about three-fourths as wide as first 
submarginal cell, emitting radius at middle; mar- 
ginal cell longer beyond than beneath stigma, 
with metacarpus somewhat less than twice as 
long as apical truncation. Pygidial plate with 
moderately wide rounded apex. Tarsal claws 
simple. 

Frons and vertex minutely tessellate, rather 
dull, and with very fine close punctures on frons 
and sides of face below antennae; punctures of 
clypeus coarser and moderately close. Meso- 
scutum delicately tessellate, shining, with fine 
punctures widely separated on posterior half, and 
moderately close anteriorly and on scutellum. 
Mesopleura minutely tessellate, with fine sepa- 
rated punctures. Abdomen dullish, minutely 
tessellate and with fine distinct punctures. Pu- 
bescence white, that on face short, erect, and 
moderately dense, and that on cheeks and thorax 
longer; hair of scutum thin, except on anterior 
border, and more than twice as long as diameter 
of ocelli. Apical fimbria of abdomen rather long 
and dense. Scopal hair of hind tibiae fine and 
simple and not greatly longer than width of tibia. 

Length, about 5 mm; anterior wing, 3 mm; 
width of abdomen, 1.5 mm. 

HoLotyPE FEMALE.—Martinez, Contra Costa County, 
California, 11 Sep. 1966 (R. M. Brown). 

PARATYPE.—One female, same data as type. 


Types in collection of the California Academy of Sciences, 
San Francisco. 


Perdita infuscata, new species 


This is the fifth Hexaperdita recorded from 
Baja California. The female runs to foveata Tim- 
berlake in the key but differs in the dusky wings 
with dark nervures, and more nearly polished 
mesoscutum with scattered fine punctures, and 
in the shorter marginal cell, as well as in the lack 
of light markings. 

FEMALE.—Head and thorax dark green and 
abdomen black, without light markings except 
tubercles white, and anterior margin of pronotum 


284 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 10 


very narrowly whitish. Legs dark, with anterior 
side of front tibiae yellowish. Scape and pedicel 
of antennae black, flagellum more fuscous. 
Tegulae dark testaceous. Wings dusky hyaline, 
nervures and margins of stigma fuscous. 

Head somewhat broader than long; cheeks 
strongly receding, about one-third as wide as 
eyes. Posterior ocelli very slightly closer to each 
other than to margin of eyes or occipital margin. 
Antennae inserted below middle of face, with 
flagellum moderately clavate and joints mostly 
about as wide as long. Facial foveae oval, very 
close to margin of eyes, about three times longer 
than wide and about half as long as space between 
level of antennae and anterior ocellus. Face 
below level of antennae gently convex, with disk 
of clypeus broader than high and rounded at 
summit. Mandibles and proboscis concealed in 
holotype. Pterostigma as long and as wide as first 
submarginal cell; marginal cell equal beneath and 
beyond stigma with metacarpus about one-fourth 
longer than apical truncation. Pygidial plate con- 
cealed in holotype, apparently rather narrowly 
rounded at apex. 

Frons very minutely granular tessellate, dull 
and minutely punctured to level of ocelli. Face 
below level of antennae more shining and rather 
closely and finely punctured. Mesoscutum 
obscurely tessellate, shining and with scattered 
fine punctures; scutellum with coarser dense 
punctures, and mesopleura shining, with sepa- 
rated punctures. 

Pubescence fine, short and thin on face and 
mesoscutum, and denser and longer on cheeks. 
Abdomen depressed, short, widest on third seg- 
ment, with short, thin hair fringing lateral margin 
of tergites 2 to 4, and apical fimbria on tergites 5 
and 6 moderately dense and long. Hind tibiae 
narrow, with scopal hair fine and nearly twice as 
long as width of tibia. 

Length, 4.5 mm; anterior wing, 3.0 mm; width 
of abdomen, 1.5 mm. 

HoLotTyPE FEMALE.—17.6 km (11 miles) W of Santo To- 
mas, Baja California Norte, Mexico, 137 m(450 ft), 11 Jul. 1969 
(S. C. Williams and V. F. Lee). 


Type in collection of the California Academy of Sciences, 
San Francisco. 


Perdita zavortinki, new species 
(Figures 1 and 2.) 
This little species of Hexaperdita is closely 


allied to heterothecae Cockerell, with the clyp- 
eus mainly black, the lateral marks triangular and 


reaching facial foveae, and abdomen black with 
small transverse white mark on each side of the 
third tergite. In the male the clypeus and trans- 
verse lateral marks are white and the abdomen 
dark. 

FEMALE.—Head and thorax dark blue-green, 
clypeus and supraclypeal area black, with lateral 
marks triangular and intruding acutely between 
foveae and margin of eyes, and small contiguous 
spot on each side of clypeus mainly on lateral 
extensions white. Thorax dark except tubercles 
white. Abdomen black, with small transverse 
white mark on each side of tergite 3. Pygidial 
plate testaceous. Legs black, extreme apex of 
femora, anterior side of front and middle tibiae 
and their tarsi yellowish white. Antennae fus- 
cous, scape white beneath, and flagellum yel- 
lowish brown beneath. Labrum and mandibles 
testaceous, latter white at base. Tegulae testa- 
ceous. Wings somewhat whitish hyaline, ner- 
vures testaceous, margins of stigma and subcosta 
more brownish. 

Head broader than long; cheeks slightly reced- 
ing, about two-thirds as wide as eyes. Posterior 
ocelli approximately their distance apart from 
margin of eyes and occipital margin. Facial 
foveae strongly impressed, about half their own 
width from margin of eyes and reaching from 
level of antennal sockets about two thirds of dis- 
tance to level of anterior ocellus. Antennae 
slightly clavate with middle joints of flagellum as 
long as thick. Disk of clypeus gently convex, 
much broader than high, and rounded on summit. 
Proboscis moderately long, galeae about equal to 
stipites. Pterostigma three-fourths as wide as first 
submarginal cell; marginal cell slightly shorter 
beneath than beyond stigma, with metacarpus 
one-half longer again than apical truncation. Py- 
gidial plate narrow at apex. 

Head and thorax shining, almost impunctate, 
frons finely tessellate and mesonotum polished. 
Pubescence whitish, fine, erect and thin, with 
face and mesonotum nearly bare. 

Length, 4 mm; anterior wing, 2.9 mm; width of 
abdomen, 0.9 mm. 

MALE.—Head and thorax dark blue-green, 
mandibles, with exception of refotestaceous api- 
cal third, white; clypeus and transverse lateral 
marks reaching no higher than summit of 
clypeus, white. Labrum more tinged with tes- 
taceous. Thorax entirely dark. Abdomen black- 
ish without markings. Legs dark, apex of fem- 
ora, anterior side of front tibiae, bases of middle 


TIMBERLAKE: NEW PERDITA 


and hind tibiae on anterior sides, and all tarsi 
white. Antennae dark, scape white beneath. 
Tegulae testaceous. Wings whitish hyaline, ner- 
vures pallid. 

Head much broader than long; cheeks strongly 
receding, nearly as broad as eyes. Occipital con- 
cavity, as seen from above and behind, shallow 
and about one-half as wide as greatest width of 
head at level of upper part of eyes. Posterior 
ocelli about their distance apart from margin of 
eyes and occipital margin. Facial foveae linear, 
about three times longer than wide. Antennae 
inserted much below middle of face, antennal 
sockets about their distance apart from margin of 
eyes. Disk of clypeus broader than high and 
broadly rounded at summit. Proboscis and wing 
venation as in female. 

Head and thorax smooth and shining, anterior 
half of frons with very minute close punctures, 
mostly absent on upper half. Vertex minutely 
tessellate, duller than frons, and impunctate. 
Mesoscutum delicately and minutely tessellate, 
with fine, sparse punctures, and mesopleura simi- 
lar but more closely punctured. Pubescence 
longer and denser than in female, erect and mod- 
erately long on cheeks and sides of thorax, and 
thinner on face, and mesonotum. Hair of legs 
longest and densest on front femora, and hair of 
hind tibiae fine and somewhat longer than 
greatest width of tibia. Abdomen depressed, with 
very short fine hair on apical segments. Subgeni- 
tal plate very similar to that of heterothecae 
(Timberlake 1956: Fig. 298), and genital armature 
as figured here. 

Length, about 3.5 mm; anterior wing, 2.9 mm; 
width of abdomen, 0.8 mm. 

HOLOTYPE FEMALE AND ALLOTYPE MALE.—0.8 km (0.5 
miles) NE of Apache, Cochise County, Arizona, resting on 
ground, 30 Aug. 1972 (T. J. Zavortink). 


Types in collection of the California Academy of Sciences, 
San Francisco. 


Subgenus Perditella Cockerell 
Perdita pusillissima, new species 


This is a new species of Perditella, allied to P. 
minima Cockerell and similar in color but marked 
with black. As there is only one male, dissection 
of the terminalia for illustration was not at- 
tempted. 

FEMALE.—General color testaceous, similar 
to that of minima but with a subquadrate black 
mark on each side of posterior part of mesoster- 
num. In paratype, large mark on frons, vertex 


285 


and anterior part of mesoscutum also black. An- 
tennae and legs testaceous, but flagellum dusky, 
more so in paratype. Tegulae testaceous. Wings 
whitish hyaline, nervures pallid. 

Head rotund, as broad as long; cheeks moder- 
ately receding, about half as wide as eyes. Facial 
foveae hardly more than twice as long as wide, 
and somewhat wider than interval between them 
and margin of eyes. Antennae inserted below 
middle of face, scape slender, not reaching ocelli, 
flagellum slightly clavate, with middle joints 
about as long as thick. Face below level of anten- 
nae gently convex. Mandibles tapering, acute. 
Proboscis moderately elongate, galeae about 
equal to stipites. Pterostigma large, wider than 
first submarginal cell, evenly rounded within; 
marginal cell in large part beneath stigma, with 
metacarpus about half as long as apical trunca- 
tion. Second submarginal cell narrowed above, 
with nervures meeting or almost meeting where 
joining radius, the second strongly curved. 

Head and thorax shining, smooth and im- 
punctate, with very short, thin, erect pubescence 
on cheeks, occiput, anterior part of scutum, and 
underpart of thorax. Scopal hair of hind legs thin 
and only slightly longer than greatest width of 
tibia. 

Length, 3.75 mm; anterior wing, 2.4 mm; width 
of abdomen, 6.8 mm. 

MALE.—Testaceous, with vertex, mesonotum 
and mark on each side of posterior part of meso- 
sternum black. Antennae and legs testaceous. 
Wings whitish hyaline, with pallid nervures. 

Head subquadrate, as long as wide, distinctly 
wider than thorax, broadened in front, with inner 
orbits diverging anteriorly; cheeks nearly as 
wide anteriorly as eyes but much wider behind, 
and with small triangular laminate process an- 
teriorly. Eyes rather small, on anterior two thirds 
of head. Occiput with semicircular emargination, 
about half as wide as width of head at summit of 
eyes. Posterior ocelli somewhat less than twice 
their distance apart from margin of eyes and oc- 
cipital margins. Facial foveae punctiform, close 
to margin of eyes, at two thirds of length of eyes 
from anterior end. Antennae inserted well below 
middle of face, scape slender, flagellum weakly 
clavate, reaching level of base of wings, with 
joints about as long as thick. Clypeus convex, 
prominent, rounded above, with two porrect 
processes on anterior border, about half as long 
as interval between their bases. Mandibles gently 
and convexly curved, tapering to very acute apex 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 10 


Zavortinki 


©) 
medialis 


Figures 1-4. Dorsal and lateral views of male genitalia of 
Perdita. Figs. 1-2, P. zavortinki; Figs. 3-4, P. medialis. 


and reaching to base of each other. Proboscis 
and vertex as in female. 

Head and thorax shining, impunctate, with ex- 
tremely minute and delicate tessellation on face. 
Pubescence very thin and short, with face, 
cheeks and mesonotum bare. 

Length, 3.2 mm; anterior wing, 2.3 mm; width 
of abdomen, 0.6 mm. 


HOLOTYPE FEMALE AND ALLOTYPE MALE.—1.6 km (1 mile) 


NE of Portal, Cochise County, Arizona, on Baccharis , 3 Aug. 
1972 (T. J. Zavortink). 

PARATYPE.—One female taken with types. 

Types in collection of the California Academy of Sciences, 
San Francisco. 


Subgenus Perdita Smith 
OCTOMACULATA GROUP 
Perdita esmeraldensis, new species 


This species falls in the keys (Timberlake 1960: 
10; 1968: 65) near ensenadensis Timberlake and 
sejuncta Timberlake. It is most similar to en- 
senadensis but differs in the pattern of abdominal 
markings and in having the pygidial plate finely 
notched at apex. 

FEMALE.—Head and thorax dark green, la- 
brum and dark part of face below level of anten- 
nae black. Mandibles at base creamy white, 
shading through testaceous to red at apex; me- 
dian oval mark on clypeus pointed above, and lat- 
eral marks, filling space between margin of eyes 
and clypeus, creamy white, remaining anterior 
border of face dark (rather broadly) on each side. 
Tubercles white, but pronotum otherwise dark. 
Abdomen black, with white bands on tergites 1 
to 5, reduced to well separated marks on tergite 
1, interrupted narrowly on tergites 2, 3 and 5, but 
entire on 4; touching full width of lateral foveae 
and narrowed medially on tergite 2, ending close 
to lateral margin of segments on tergites 3 and 4, 
with that on tergite 5 more enclosed. Venter dark, 
but sternite 6 and pygidial plate testaceous and 
sternite 5 tinged with testaceous apically. Legs 
black, with extreme apex of front and middle 
femora, anterior side of their tibiae and tarsi, and 
hind knees white. Scape of antennae black, 
flagellum fuscous, becoming yellowish brown 
beneath. Tegulae testaceous. Wings somewhat 
opaquely subhyaline, nervures and margins of 
stigma testaceous. 

Head somewhat broader than long; cheeks 
rounded, moderately receding and half as wide as 
eyes. Posterior ocelli about their distance apart 
from occipital margin and somewhat more distant 
from margin of eyes. Facial foveae slender, about 
half their width from margin of eyes and reaching 
about halfway from level of antennal sockets to 
level of anterior ocellus. Antenna inserted below 
middle of face, moderately clavate, with middle 
joints of flagellum somewhat thicker than long. 
Face below level of antennae convex from side to 
side, with disk of clypeus considerably broader 
than high and rounded at summit. Mandibles ta- 


TIMBERLAKE: NEW PERDITA 


pering and acute. Proboscis retracted in holotype 
and slightly exceeding proboscidial fossa. Ptero- 
stigma half as wide as first submarginal cell; mar- 
ginal cell longer beyond than beneath stigma, 
with metacarpus about one-third longer again 
than apical truncation. Pygidial plate with small 
notch at apex. 

Frons and vertex very minutely granular, dull 
and impunctate, with face below level of anten- 
nae shining and obscurely punctured. Mesoscu- 
tum like frons, but slightly less minutely granu- 
lar tessellate. Mesopleura and propodeum very 
minutely tessellate and shining. Pubescence fine, 
erect, whitish, rather dense and about twice as 
long as diameter of ocelli on cheeks, slightly 
shorter and less dense on mesonotum, and longer 
on underside of thorax. Apical fimbria of abdo- 
men long and thin. Scopal hair of hind legs fine, 
obliquely inclined and somewhat longer than 
greatest width of tibia. 

Length, 5 mm; anterior wing, 2.6 mm; width of 
abdomen, 1.3 mm. 

HOLOTYPE FEMALE.—Lida, at summit, 2255 m (7400 ft), 
Esmeralda County, Nevada, 15 Jul. 1966 (P. H. Arnaud, Jr.). 


Type in collection of the California Academy of Sciences, 
San Francisco. 


Perdita inyoensis, new species 


This is a mainly dark species from the Angus 
Mountains. The specimens are in rather poor 
condition for study; they appear to agree best 
with the Octomaculata Group. In the key (Tim- 
berlake 1960: 4), it runs out at polygonellae Tim- 
berlake but the abdomen is black with the middle 
segments translucent reddish. 

FEMALE.—Head and thorax very dark green, 
with dark part of face below level of antennae and 
mesonotum black. Mandibles, with exception of 
reddish tips, creamy white; median stripe on 
clypeus, triangular lateral marks (broad in front, 
about twice as long as wide and ending acutely at 
anterior end of foveae, with slight extension be- 
tween foveae and margin of eyes), tubercles and 
small spot on each side of hind margin of pro- 
notum creamy white. Abdomen black, with mid- 
dle segments translucent reddish, and with faint 
nubilous whitish mark on disk of tergite 1 extend- 
ing forward onto basal declivity. Antennae and 
legs dark, but apex of front femora, anterior side 
of front tibiae and front tarsi pale yellow. Wings 
somewhat whitish hyaline, nervures testaceous. 
Tegulae testaceous. 

Head about as long as wide, with cheeks half as 


287 


wide as eyes and receding. Posterior ocelli 
somewhat closer to margin of eyes and occipital 
margin than their distance apart. Facial foveae 
well impressed, about four times longer than 
wide, close to margin of eyes, and reaching from 
lower level of antennal sockets about three 
fourths of distance to level of anterior ocellus. 
Antennae mainly missing, but scape slender, of 
normal length. Face below level of antennae 
gently convex from side to side, with disk of 
clypeus as broad as high and rounded at summit. 
Mandibles tapering and acute. Proboscis moder- 
ately long, galeae shorter than stipites. Ptero- 
stigma three-fourths as wide as first submarginal 
cell; marginal cell slightly longer beneath than 
beyond stigma, with apical truncation and 
metacarpus equal. Pygidial plate nearly con- 
cealed in holotype, but very narrow at apex. 

Head and thorax dullish, minutely tessellate, 
with minute punctures between foveae and an- 
tennal sockets and a few minute punctures on 
anterior border of mesoscutum. Face below level 
of antennae polished and impunctate. Pubes- 
cence whitish, fine, thin and erect, with only a 
few hairs on anterior border of scutum, but front 
coxae with dense white hair. 

Length, about 5 mm; anterior wing, 3.5 mm; 
width of abdomen, 1.3 mm. 

HOLOTYPE FEMALE.—Angus Mountains, Inyo County, 
California, 12 May 1948 (E. R. Leach). 

PARATYPE.—One female taken with the type. 


Types in collection of the California Academy of Sciences, 
San Francisco. 


Perdita medialis, new species 
(Figures 3 and 4.) 


The female of medialis has the face below level 
of antennae white, with a large bilobed dark 
mark, partly on the clypeus. In the key (Timber- 
lake 1960: 10, 19) to the Octomaculata Group, 
medialis runs to swenki Crawford but differs by 
the black mark on the face, by the light pronotum 
with dark band from flank to flank, and by the 
pale yellow legs. 

FEMALE.—Head and thorax dark green, man- 
dibles, with exception of reddish apical third, 
creamy white; labrum and face below level of 
antennae creamy white, lateral marks ending 
broadly at level of antennae. Large mark on each 
side of upper half of clypeus (interval nearly as 
wide as marks and moderately narrowed above) 
and supraclypeal area intruding roundly between 
antennae black, with subantennal plates slightly 


greenish. Pronotum white, with green band from 
flank to flank. Abdomen pale yellow, with small 
spot on lateral margins of tergite 1 at summit of 
basal declivity, and narrow band at junction of 
tergites 1-2 to 4-5 black, with that portion of 
bands at apex of segments obliquely narrowed at 
outer ends, and part at base of segments pro- 
duced into short curved spur at outer ends on 
tergites 3 and 4, and that on tergite 2 confluent at 
outer ends with black lateral foveae. Tergite 6 
testaceous. Legs pale yellow, with hind coxae 
except at apex, hind tibiae except at base, and 
hind tarsi fuscous, and middle tibiae dusky be- 
hind. Antennae fuscous, scape broadly pale yel- 
low beneath and flagellum dull yellowish be- 
neath. Tegulae whitish, with subhyaline spot on 
outer margin. Wings whitish hyaline with pale 
testaceous nervures and stigma. 

Head slightly longer than wide, cheeks reced- 
ing, about half as wide as eyes. Posterior ocelli 
closer to margin of eyes and occipital margin than 
their distance apart. Facial foveae very slender, 
close to margin of eyes, and reaching about two 
thirds of distance from level of antennal sockets 
to level of anterior ocellus. Antennae normal for 
group, with middle joints of flagellum as long as 
wide. Face below level of antennae gently con- 
vex, clypeus rounded at summit, with lateral ex- 
tensions broadly inflexed. Mandibles tapering 
and acute. Proboscis moderately elongate, galeae 
about equal to stipites. Pterostigma hardly more 
than half-as wide as first submarginal cell; mar- 
ginal cell longer beyond than beneath stigma, 
with metacarpus nearly twice as long as apical 
truncation. Pygidial plate about as long as wide 
at base, with slightly crenate sides converging to 
rounded and slightly retuse apex. 

Frons and vertex very minutely granular, matt 
and obscurely punctate. Face below level of 
antennae smooth, shining and impunctate. 
Mesonotum minutely tessellate, densely and 
very finely punctured and more shining than 
frons. Mesopleura shining, minutely tessellate 
and with close minute punctures. Propodeum 
tessellate and shining with minute dense 
punctures on lateral ridges. Pubescence whitish, 
fine, erect, rather long on cheeks and sides of 
thorax, and short, erect on anterior part of frons, 
and short, erect and yellowish and rather dense 
on scutum and scutellum, but face below anten- 
nae bare. Apical fimbria of abdomen long and 
rather dense. Scopal hair of hind tibiae normal for 
group. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 10 


Length, 6 mm; anterior wing, 4 mm; width of 
abdomen, 1.8 mm. 

MALE.—Head and thorax dark green, marked 
with yellow, probably pale, but reddened by 
cyanide in allotype. Mandibles, except tips, la- 
brum and face to level of foveae yellow; under- 
side of head, hypostoma, walls of proboscidial 
fossa and very narrow postorbital orbits to mid- 
dle of eyes also light. Pronotum mainly yellow, 
including tubercles, but with broad dark band de- 
scending on flank, with posterior spur at outer 
ends protruding narrowly to inferior margin of 
tubercles. Abdomen yellow with base of tergite 1 
and band at junction of segments black; first two 
bands narrow but wider on following segments 
with yellow on tergite 5 enclosed; tergite 6 black, 
with very narrow, abbreviated and interrupted 
yellow band; tergite 7 also black, yellow on apical 
lobe, and venter uniformly yellow. Legs yellow, 
narrow line behind on inferior margin of front 
femora and tibiae, broader marks behind on mid- 
dle femora and tibiae, hind femora beneath, hind 
tibiae (except in front) and hind tarsi fuscous. 
Antennae yellow, pedicel black above and flagel- 
lum dusky. Tegulae testaceous. Wings sub- 
hyaline, nervures testaceous. 

Head somewhat broader than long; cheeks but 
little receding, rounded and three-fourths as wide 
as eyes. Posterior ocelli slightly more than their 
distance apart from margin of eyes and occipital 
margins. Antennae inserted slightly below mid- 
dle of face, scape about four times longer than 
thick, flagellum slightly clavate, with middle 
joints as long as thick. Face below level of anten- 
nae gently convex from side to side, with disk of 
clypeus broader than high and truncate at sum- 
mit. Mandibles acute and reaching somewhat be- 
yond far margin of labrum. Proboscis briefly ex- 
ceeding proboscidial fossa. Venation as in 
female. 

Frons minutely granular, dull and impunctate. 
Face below level of antennae moderately shining 
and impunctate. Mesonotum dull, very minutely 
tessellate and obscurely punctured. Mesopleura 
shining and minutely punctured. Abdomen elon- 
gate oval, depressed, with apical tergite nar- 
rowly rounded at apex. Pubescence whitish, fine, 
moderately long, and of usual density on cheeks 
and underside of thorax, very short and thin on 
face, and thin, erect and short on mesoscutum. 
Terminalia as figured. 

Length, 6 mm; anterior wing, 4 mm; width of 
abdomen, 1.5 mm. 


TIMBERLAKE: NEW PERDITA 


HOLOTYPE FEMALE AND ALLOTYPE MALE.—Gran Quivira, 
Torrance County, New Mexico, 1,981 m (6500 ft), 18-20 Aug. 
1967 (H. B. Leach). 

PARATYPES.—1wo females, same data as for types. 

Types in collection of the California Academy of Sciences, 
San Francisco, except one paratype in collection of the Univer- 
sity of California, Riverside. 


Perdita pauliana, new species 
(Figures 5, 6, 7.) 


This species is closely allied to swenki Craw- 
ford and aridella Timberlake, near which it runs 
in the key (Timberlake 1960: 19, 20) to the Oc- 
tomaculata Group. It differs in having the clypeus 
mainly yellow, and the light bands about equal to 
the dark intervals, and enclosed on tergites | to 5, 
with last two segments nearly all yellow. The 
head in the type-specimen is moderately enlarged 
and subquadrate. 

MALE.—Head and thorax dark green. Gular 
region, inferior orbits on anterior fourth of eyes, 
mandibles except red tips, labrum and anterior 
half of face to level of foveae, clear yellow. 
Thorax dark, except collar narrowly; transverse 
mark on each side of hind margin of pronotum, 
and small spot on tubercles yellow. Abdomen 
yellow, with band at base of tergite | and band at 
junction of segments 1-2 to 5—6 fuscous, with 
light bands equal to dark intervals and almost 
reaching lateral margins, very uniform in width 
with median notch behind on tergites | to 4. Dark 
banding on two apical segments subobsolete. 
Front legs yellow, with femora black behind ex- 
cept at apex; middle legs dark, with apex of fem- 
ora, tibiae and tarsi yellow, and hind legs dark 
with knees yellow. Antennae yellow, spot at apex 
of scape above and pedicel black, and flagellum 
brownish above. Tegulae fuscous. Wings 
hyaline, nervures testaceous, margins of stigma 
and subcosta brownish. 

Head moderately enlarged, subquadrate, 
broader than long and broader than thorax. 
Cheeks slightly receding, about as wide as eyes. 
Posterior ocelli about their distance apart from 
occipital margin but closer to margin of eyes. 
Facial foveae small, twice as long as wide, 
situated close to margin of eyes and somewhat 
below halfway point between levels of antennal 
sockets and anterior ocellus. Antennae inserted 
below middle of face, moderately clavate, with 
middle joints of flagellum as long as thick. Face 
below level of antennae rather strongly convex 
with disk of clypeus as broad as high, nearly 
truncate at summit, and lateral extensions 


289 


broadly inflexed and only partly visible in frontal 
aspect. Mandibles simple and reaching far margin 
of labrum. Proboscis retracted in holotype and 
somewhat exceeding proboscidial fossa. Ptero- 
stigma about half as wide as first submarginal 
cell; marginal cell about equal beneath and be- 
yond stigma, with metacarpus distinctly longer 
than apical truncation. 

Frons and vertex very dull and impunctate, 
and face below antennae shining with faint, 
sparse punctures. Mesonotum tessellate, dull, 
but more shining than frons, and with fine, mod- 
erately close, obscure punctures. Mesopleura 
more shining than notum, finely and sparsely 
punctured. Pubescence white, rather short, erect 
and dense on cheeks, very short and thin on face 
above antennae, and short, erect and thin on 
mesonotum. 

Abdomen depressed, with apical segment pro- 
vided medially with narrow obtuse lobes. Sub- 
genital plate and genital armature as figured 
(Figs. 5-7). 

Length, about 4.5 mm; anterior wing, 3.0 mm; 
width of abdomen, 1.6 mm. 

HOLOTYPE MALE.—Lida, at summit, Esmeralda County, 
Nevada, 2255 m (7400 ft) 15 Jul. 1966 (P. H. Arnaud, Jr.). 


Type in collection of the California Academy of Sciences, 
San Francisco. 


Perdita willcoxiana, new species 


This species of the Octomaculata Group runs 
to the three species separated in couplets 73-74 of 
the key (Timberlake 1960: 9-10), and agrees best 
with cazieri Timberlake, but differs in having the 
clypeus mainly black, with a median stripe, and 
the lateral marks small and transverse. 

FEMALE.—Head and thorax dark green, more 
bluish on propodeum, with dark part of clypeus 
and supraclypeal area black. Median stripe on 
clypeus, dilated anteriorly, very narrow lateral 
margin of disk and small lateral mark between 
clypeus and eyes creamy white. Tubercles and in 
some cases small spot on each side of hind margin 
of pronotum also white. Abdomen black with 
narrow white band on tergites | to 4, narrowly 
interrupted medially, curved backward at outer 
ends and reaching lateral margins on tergites 2 
and 3, somewhat abbreviated on tergite 4 and that 
on tergite 1 in some cases divided into four trans- 
verse marks. Legs black, extreme apex of front 
femora, anterior side of front and middle tibiae 
and front tarsi pale yellow. Mandibles white ex- 
cept red at apex. Antennae dark, but radicle joint 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 10 


apicalis 


10 


gracilior 


bifasciata 


FiGures 5—16. Dorsal and lateral views of male genitalia, and subgenital plate of Perdita. Figs. 5-7, P. pauliana; Figs. 8-10, P. 


gracilior; Figs. 11-13, P. apicalis; Figs. 14-16, P. bifasciata. 


of scape white, and flagellum yellowish beneath. 
Tegulae testaceous. Wings slightly whitish 
hyaline, nervures testaceous, margins of stigma 
and subcosta more brownish. Pygidial plate tes- 
taceous. 

Head as broad as long, cheeks receding, about 
half as wide as eyes. Posterior ocelli somewhat 
closer to margin of eyes and occipital margin than 
their distance apart. Facial foveae narrow, close 
to margin of eyes and reaching about halfway 
from level of antennal sockets to level of anterior 
ocellus. Antennae normal for group, with middle 
joints of flagellum about as long as thick. Face 
below level of antennae gently convex, with disk 
of clypeus somewhat broader than high and sub- 
truncate at summit. Mandibles simple and acute. 
Proboscis moderately elongate, with galeae equal 
to stipites. Pterostigma hardly more than half as 
wide as first submarginal cell; marginal cell about 
as long as stigma, with parts beneath and beyond 
stigma equal, and metacarpus distinctly longer 


than apical truncation. Abdomen depressed, 
about three times longer than wide, with pygidial 
plate notched at apex. 

Frons and vertex very minutely granular, dull, 
and impunctate except close to base of anten- 
nae. Face below level of antennae more shining, 
with shallow obscure punctures on clypeus. 
Mesonotum minutely tessellate, more shining 
than frons, and with minute close punctures. 
Pubescence white, short, erect, thin on face and 
abundant on mesonotum. Scopal hair of hind 
tibiae distinctly longer than greatest width of 
tibia. 

Length, about 6 mm; anterior wing, 3.7 mm; 
width of abdomen, 1.5 mm. 


HOLOTYPE FEMALE.—Willcox Dry Lake, Cochise County, 
Arizona, 17 Sep. 1969 (B. A. Tilden). 

PARATYPE.—Three females taken with the holotype. 

Types in collection of the California Academy of Sciences, 
San Francisco, except one paratype in collection of the Univer- 
sity of California, Riverside. 


TIMBERLAKE: NEW PERDITA 


VENTRALIS GROUP 


Perdita diminutiva, new species 


This is an isolated diminutive member of the 
Ventralis Group, difficult to place in any particu- 
lar section of the group, but the concave sides of 
the face are suggestive of the claypolei section. In 
the key to the group (Timberlake 1962: 13) it 
probably runs best to couplet 100 where it can be 
distinguished by the yellow color of its face below 
level of antennae and by the dark thorax and 
abdomen. The minute terminalia were lost in dis- 
section, but the genital armature was similar to 
that in Figure 771 (op. cit.), and I believe that the 
subgenital plate was wide at apex and notched. 

MALE.—Head and thorax dark green. Mandi- 
bles, with exception of reddened extreme apices, 
yellow; labrum and face below level of antennae, 
yellow, with very broad parocular parts extend- 
ing bluntly to level of foveae. Thorax dark, ex- 
cept tubercles yellow, and abdomen dark without 
markings. Legs dark, except front tibiae and tarsi 
yellow. Antennae yellow, with flagellum fuscous 
above, but streaked with yellow at juncture of 

joints. Tegulae dark testaceous. Wings dusky 
hyaline, nervures fuscous. 

Head subquadrate, as broad as long, and 
broader than thorax. Cheeks very broad, reced- 
ing slightly, rounded, but narrowed anteriorly 
and not dentate. Posterior ocelli about their dis- 
tance apart from occipital margin and twice as far 
from margin of eyes. Facial foveae very obscure. 
Antennae inserted just below middle of face, and 
reaching level of anterior ocellus at juncture of 
second and third flagellar joints; scape very 
thick, only slightly longer than wide; flagellum 
thick, cylindrical, except narrowed slightly at 
base, with joints mainly thicker than long. Su- 
praclypeal area and subantennal plates mainly 
equal in size and together equal to parocular area 
on each side of face, with median part ridged and 
parocular areas shallowly concave. Disk of 
clypeus about twice as broad as high, with lateral 
extension gradually narrowed to base of mandi- 
bles. Labrum twice as broad as long. Mandibles 
tapering from base to acute apex and reaching far 
margin of labrum. Proboscis short, galeae shorter 
than stipites. Pterostigma about half as wide as 
first submarginal cell; marginal cell nearly equal 
beneath and beyond stigma, with metacarpus 
slightly longer than apical truncation. 

Frons and vertex very minutely granular tes- 
sellate, obscurely punctured and dull; yellow part 


291 


of face shining and impunctate. Mesonotum deli- 
cately tessellate, shining and with widely scat- 
tered minute punctures. Mesopleura shining, tes- 
sellate and sparsely punctured. Pubescence 
whitish, short, erect, fine and thin, most evident 
on cheeks, sides of thorax and anterior border of 
scutum, with face bare. Hind tibiae increasing 
gradually in width from narrow base to apex, with 
fine sparse, semi-erect hairs about equaling 
greatest width. 

Length, 5 mm; anterior wing, 2 mm; width of 
abdomen and head, 0.9 mm. 

HOLOTYPE MALE.—Agua Caliente (San Carlos), 18.5 km 
(11.5 miles) E of Maneadero, Baja California, Mexico, 6 Jul. 
1973 (P. H. Arnaud, Jr.). 

Type in collection of the California Academy of Sciences, 
San Francisco. 


Perdita gracilior, new species 
(Figures 8, 9, 10.) 


This little species from Baja California evi- 
dently belongs in the Ventralis Group and prob- 
ably is as closely related to exilis Timberlake as to 
any other species, but differs greatly in sparcity of 
markings. In the key to group (Timberlake 1962: 
2-14), the male runs near semicrocea Cockerell 
and the female runs with some discrepancy near 
exilis but is entirely dark blue or black without 
markings. 

FEMALE.—Head and thorax very dark blue, 
clypeus, supraclypeal area and mesonotum 
black; abdomen black and testaceous beneath; 
pygidial plate ferruginous. Light marking entirely 
absent. Legs dark, with apex of front femora, 
front tibiae and all tarsi yellowish white. Anten- 
nae dark, flagellum brownish but whitish be- 
neath. Tegulae pale testaceous. Wings whitish 
hyaline, nervures pallid, stigma yellowish, its 
margins and apical half of costal nervure infus- 
cated. Mandibles yellowish white, red at apex. 

Head slightly broader than long, with cheeks 
strongly receding, somewhat more than half as 
wide as eyes. Lateral ocelli about their distance 
apart from margin of eyes and from occipital 
margin. Facial foveae well impressed, about 
three times as long as wide and close to margin of 
eyes, and about half as long as space between 
antennal sockets and anterior ocellus. Antennae 
inserted just below middle of face, with scape 
reaching hardly more than halfway to level of 
anterior ocellus; flagellum moderately clavate, 
with middle joints slightly thicker than long. Face 
below level of antennae gently convex, with fa- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 10 


cial sutures indistinct. Mandibles parallel sided, 
subacute at apex. Proboscis rather short, galeae 
shorter than stipites. Pterostigma about as long 
and wide as first submarginal cell; marginal cell 
much longer beneath than beyond stigma, with 
metacarpus shorter than apical truncation. Py- 
gidial plate longer than wide with sides con- 
verging to narrowly rounded apex; disk flat. 

Head and thorax shining; face below level of 
antennae and mesonotum polished, mainly im- 
punctate; frons very minutely tessellate, with 
minute obscure punctures in area between foveae 
and antennal sockets. Pubescence whitish, fine 
and short, few short erect hairs visible on margins 
of scutum. Apical fimbria rather long but thin. 
Scopal hair of hind legs about as long as greatest 
width of tibia. 

Length, 5 mm; anterior wing, 3.6 mm; width of 
abdomen, 1.5 mm. 

MALE.—Head and thorax very dark blue, with 
mesonotum black. Anterior margin of clypeus 
mainly white, and mandibles white, but reddened 
at apex. Abdomen black, apical segment and ven- 
ter testaceous, but abdomen sometimes all tes- 
taceous except basal tergite, or apical segment 
more ferruginous. Legs black, apex of front fem- 
ora, front tibiae and all tarsi yellowish white. 
Scape black, narrowly yellowish beneath, flagel- 
lum fuscous and yellowish brown beneath. Wings 
whitish hyaline, nervures pallid. 

Head rotund, slightly broader than long; 
cheeks strongly receding, about half as wide as 
eyes and with small anterior conical process 
about as long as wide at base. Lateral ocelli about 
their distance apart from margin of eyes and ap- 
proximately same distance from occipital mar- 
gin. Facial foveae punctiform. Face depressed, 
slightly ridged between antennae and very 
slightly convex from side to side below level of 
antennae. Mandibles very slender, evenly 
curved, simple and nearly reaching to base of 
each other. Proboscis moderately long, galeae 
about as long as stipites. Pterostigma and vena- 
tion as in female. 

Head and thorax shining, virtually impunctate, 
but frons minutely tessellate and slightly dullish. 
Pubescence whitish, very fine, thin and short. A 
few hairs on margin of mesoscutum, but face 
nearly bare. 

Length, 3 mm; anterior wing, 2.9 mm; width of 
abdomen, 1.2 mm. 


HOLOTYPE FEMALE AND ALLOTYPE.—41.8 km (26 miles) S 
of San Felipe, Baja California Norte, Mexico, 15 Apr. 1965 (D. 


Q. Cavagnaro, C. E. and E. S. Ross, and V. L. Vesterby). 

PARATYPES.—Four females, 14 males, collected with 
holotype and allotype, and four damaged specimens. 

Types in collection of the California Academy of Sciences, 
San Francisco, except one pair of paratypes and two damaged 
specimens in collection of the University of California, River- 
side. 


SPHAERALCEAE GROUP 


Perdita apicalis, new species 
(Figures 11, 12, 13.) 


Although the dorsal lobes of the caulis are 
somewhat produced as in koebelei Timberlake, 
the general structural characters of both species 
are in agreement with the Sphaeralceae Group. 
In the key to this group (Timberlake 1964: 126— 
158), the male runs to couplet 188, but differs from 
cushmani Timberlake and falcata Timberlake in 
having the three apical segments of the abdomen 
testaceous, and the others mainly black, or some- 
times marked with testaceous yellow. The sub- 
genital plate (Fig. 13) of apicalis is very distinc- 
tive. The female runs in the same key (loc. cit.) to 
couplet 75, but differs from dammersi Timber- 
lake and obliqua Timberlake in having the face 
and thorax mainly dark and the abdomen banded 
with white, but testaceous at apex. 

MALE.—Head and thorax dark green, with an- 
terior corners of cheeks, mandibles except at 
apex, and face below level of antennae yellow, 
except for intrusion of green between subanten- 
nal plates and lateral marks, with the last- 
mentioned marks ending acutely. Thorax entirely 
dark. Abdomen fuscous, with narrow apical de- 
pression of tergites 2 to 4 whitish subhyaline; 
three apical segments, entire undersurface, and 
apical margin of disk of tergite 4, testaceous. 
Tergite 2 with abbreviated and interrupted white 
line at base and, in one paratype, tergites 2 and 4 
each with broad, testaceous yellow band extend- 
ing bluntly to apex of disk and narrowed on each 
side by broad, oblique, fuscous band. Legs tes- 
taceous yellow, with front and middle femora 
behind, streak behind on their tibiae, hind femora 
and tibiae except in front, and hind tarsi fuscous. 
Scape of antennae yellow, flagellum duller yel- 
low and mainly dusky above. Proboscis brownish 
testaceous, darker at base. Tegulae pale testa- 
ceous. Wings whitish hyaline, nervures pale tes- 
taceous, margins of clypeus and subcosta brown- 
ish. 

Head as broad as long, cheeks receding and 
nearly as broad posteriorly as eyes. Posterior 
ocelli about three fourths of their distance apart 


TIMBERLAKE: NEW PERDITA 


from margin of eyes and somewhat closer to oc- 
cipital margin. Antennae inserted below middle 
of face, scape reaching nearly halfway to level of 
anterior ocellus, flagellum thickly cylindrical, 
with joints as broad as long. Antennal sockets 
their distance apart from margin of eyes. Frons 
almost plane; disk of clypeus as broad as high, 
rounded at summit, with lateral extensions 
broadly inflexed and not visible in frontal as- 
pects. Extreme anterior corners of face also in- 
flexed. Mandibles very acute and reaching for 
margin of labrum. Proboscis moderately elon- 
gate; galeae reaching base of stipites in repose. 
Pterostigma as long, and about three-fourths as 
wide, as first submarginal cell; marginal cell 
longer beneath than beyond stigma, with 
metacarpus subequal to apical truncation. Tarsal 
claws bifid. 

Frons and vertex minutely tessellate, rather 
dull, and with dense minute punctures. Face 
below level of antennae shining, somewhat 
roughened by shallow punctures. Mesonotum 
polished and shining, with fine separated 
punctures. Mesopleura similarly sculptured. 
Pubescence whitish, erect, and rather dense on 
cheeks and sides of thorax, shorter on face above 
antennae, and thinner on mesoscutum. Abdomen 
about three times longer than wide, with subgeni- 
tal plate and genital armature as figured (Figs. 
11-13). 

Length, 4.5 mm; anterior wing, 3.5 mm; width 
of abdomen, 1.5 mm. 

FEMALE.—Head and thorax dark green, but 
face below level of antennae and labrum black, 
with at most small lateral marks or transverse 
lateral marks and median stripe on clypeus, 
white. Mandibles whitish, reddish on apical 
third. Thorax entirely dark. Abdomen testaceous 
beneath and on apical two segments, otherwise 
fuscous above marked with white. Tergite | with 
abbreviated and interrupted white line at summit 
of base declivity, and tergites 2 to 4 each with 
broad white band, nearly reaching lateral margins 
and almost as wide as dark intervals between 
them, with apical depression pale testaceous and 
subequal to light and dark bands. Legs dark, ex- 
treme apex of front and middle femora, together 
with anterior side of their tibiae, and tarsi yel- 
lowish white. Antennae dark, scape black, flagel- 
lum brownish and sordid yellowish beneath. 
Tegulae subhyaline testaceous. Wings whitish 
hyaline as in male. 

Head rotund, as broad as long; cheeks strongly 


293 


receding, about half as wide as eyes. Posterior 
ocelli slightly more than their distance apart from 
nearest eye and occipital margin. Facial foveae 
about four times longer than wide, very close to 
margin of eyes and reaching about three fourths 
of distance from level of antennal sockets to level 
of anterior ocellus. Flagellum of antennae slightly 
clavate and rather thick, with joints mostly about 
as broad as long. Disk of clypeus much broader 
than high and rounded at summit. Mandibles 
reaching far margin of labrum, abruptly narrowed 
on inner margin, with formation of small inner 
tooth and very acute at apex. Proboscis and 
wings as in male. Pygidial plate narrowed from 
broad base to rounded apex, with disk plane. 

Frons and vertex minutely tessellate and dull, 
with minute punctures nearly to level of ocelli; 
face below level of antennae shining, with close 
shallow punctures. Thorax polished, with 
punctures of mesonotum numerous but mostly 
well separated, and puncturation of mesopleura 
similar. Pubescence whitish, fine, erect, dense 
and long on cheeks and sides of thorax, rather 
dense on face above antennae and thin on 
mesonotum. Apical fimbria of abdomen long but 
thin. Scopal hair of hind legs more than twice as 
long as greatest width of tibia. 

Length, 5 mm; anterior wing, 3.5 mm; width of 
abdomen, 1.6 mm. 

HOLOTYPE MALE AND ALLOTYPE.—41.8 km (26 miles) S of 
San Felipe, Baja California Norte, Mexico, 15 Apr. 1965 (D. Q. 
Cavagnaro, C. E. and E. S. Ross, and V. L. Vesterby). 

PARATYPES.—One female, two males, same uata as for 
holotype and allotype. 

Types in collection of the California Academy of Sciences, 


San Francisco, except one paratype in collection of the Univer- 
sity of California, Riverside. 


Perdita bifasciata, new species 
(Figures 14, 15, 16.) 


In the key to Sphaeralceae Group (Timber- 
lake 1964: 126-158), this species runs to zebrata 
Cresson in couplet 228 and is very distinct in the 
markings and terminalia. 

MALE.—Head and thorax dark blue-green, 
mesonotum black. Mandibles light rufotesta- 
ceous; labrum, face below level of antennae with 
lateral marks extending to level of ocelli, anterior 
part of undersurface of head and posterior orbits 
to middle of eyes, yellow. Pronotum dark on 
disk, but hind margin confluent with mark on 
flanks, and tubercles yellow, but median dark 
lobe from disk descending halfway on flanks. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 10 


Prosternum yellow except lateral margins, and 
mesosternum with yellow mark on each side an- 
teriorly. Thorax otherwise dark. Abdomen fus- 
cous, with even, narrow yellow band at base of 
tergites 2 and 3, that on tergite 2 touching lateral 
foveae, other reaching lateral margins of seg- 
ment. Venter of abdomen testaceous yellow. 
Front and middle legs, apex of hind tibia, basal 
half of hind femora with entire underside yellow, 
but front femora black behind except at apex and 
front and middle tibiae dark on posterior margins. 
Antenna yellow, flagellum tinged with fuscous. 
Tegulae testaceous. Wings nearly clear hyaline, 
nervures testaceous, margins of stigma and sub- 
costa dusky. 

Head somewhat broader than long, cheeks 
moderately receding, rounded and about half as 
wide as eyes. Antennae inserted below middle of 
face, scape about four times longer than thick, 
flagellum nearly cylindrical with joints about as 
long as thick. Face convex from side to side with 
disk of clypeus broader than high and well 
rounded at summit. Mandibles tapering, acute 
and reaching far margin of labrum. Proboscis 
moderately long, galeae equal to stipites. Ptero- 
stigma four times longer than wide and hardly 


more than half as wide as first submarginal cell; ° 


marginal cell equal beneath and beyond stigma, 
with metacarpus distinctly longer than apical 
truncation. 

Head and thorax shining, minutely tessellate 
on frons, vertex and thorax, and with very minute 
and sparse punctures. Face below level of anten- 
nae polished and impunctate. Pubescence 
whitish, long, fine and moderately dense on sides 
and underside of thorax, thin on face and thin, 
short and erect on mesonotum. 

Length 5.5 mm, anterior wing 3.2 mm, width of 
abdomen 1.5 mm. 


HOLOTYPE MALE.—Southwestern Research Station, 1646 
m (5400 ft), 8 km (5 miles) W of Portal, Cochise County, 
Arizona, 11 Sep. 1965 (P. H. Arnaud, Jr.). 

PARATYPE.—One male (abdomen missing), same data as for 
holotype. 

Types in collection of California Academy of Sciences, San 
Francisco. 


Perdita perlucens, new species 


The female of perlucens is similar to lucidella 
Timberlake (1964: 204-205) in size and color, but 
has the face much more punctured, the facial 
foveae much larger, the pterostigma more slen- 


der, the marginal cell longer, with the metacarpus 
twice as long as apical truncation and the pygidial 
plate broader. 

FEMALE.—Head and thorax very dark blue, 
with clypeus and supraclypeal area black, and no 
light markings. Abdomen blackish, unmarked; 
pygidial plate testaceous. Legs dark, anterior 
side of front tibiae testaceous yellow, tarsi tes- 
taceous. Scape of antennae black, flagellum 
fuscous above and brownish beneath. Labrum 
and mandibles testaceous, with apex of mandi- 
bles darker. Tegulae testaceous. Wings slightly 
dusky hyaline, nervures and margins of stigma 
testaceous, subcosta darker. 

Head distinctly broader than long; cheeks 
strongly receding and about half as wide as eyes. 
Posterior ocelli only slightly more than their 
distance apart from margin of eyes and occipital 
margin. Facial foveae oval, twice as long as wide, 
or slightly more, close to margin of eyes and 
reaching from level of antennal sockets about two 
thirds of distance to level of anterior ocellus. 
Antennae inserted below middle of face, flagel- 
lum slightly clavate, with middle joints as wide as 
long. Disk of clypeus much broader than high, 
rounded at summit, with lateral extensions 
broadly inflexed. Mandibles simple. Proboscis 
moderate in length, not exceeding fossa in re- 
pose. Pterostigma only slightly more than half as 
wide as first submarginal cell; marginal cell as 
long beneath as beyond stigma, with metacarpus 
nearly twice as long as apical truncation. Pygidial 
plate plane with lateral margins converging to 
rounded apex. 

Head and thorax mainly polished and shining, 
but anterior half of frons minutely tessellate and, 
together with face below level of antennae, very 
closely and minutely punctured, except 
punctures of clypeus distinctly larger. Punctures 
of mesonotum sparse, but more numerous on 
anterior border. Pubescence white, rather short 
and dense on cheeks, longer on underside of 
thorax, and short and thin on face and 
mesonotum. Scopal hair of hind tibiae about 
twice as long as greatest width of tibia. 

Length, 4 mm; anterior wing, 3 mm; width of 
abdomen, 1.2 mm. 

HOLOTYPE FEMALE.—Mammoth Lakes, Mono County, 
California, 5 Aug. 1936 (Ruth and G. E. Bohart). 

PARATYPE.—Six females, taken with the holotype. 

Types in collection of the California Academy of Sciences, 


except one paratype in the collection of the University of 
California, Riverside. 


TIMBERLAKE: NEW PERDITA 295 


LITERATURE CITED 


. 1960. Ibid. Part IV. Univ. Calif. Publ. Entomol. 17(1): 


TIMBERLAKE, P.H. 1954. A revisional study of the bees of the 1-156. 
genus Perdita F. Smith, with special reference to the fauna of . 1962. Ibid. Part V. Univ. Calif. Publ. Entomol. 28(1): 
the Pacific Coast (Hymenoptera, Apoidea). Part I. Univ. 1-123. 
Calif. Publ. Entomol. 9(6): 345-432. . 1964. Ibid. Part VI. Univ. Calif. Publ. Entomol. 28(2): 
_ 1956. Ibid. Part II. Univ. Calif. Publ. Entomol. 11(5): 125-387. 
247-349. 1968. Ibid. Part VII. (Including index to parts I to 


_ 1958. Ibid. Part III. Univ. Calif. Publ. Entomol. 14(5): VII.) Univ. Calif. Publ. Entomol. 49: 1-196. 
303-410. 


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PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 11, pp. 297-305; 5 figs. 


February 8, 1978 


NEW SAND DOLLARS (ECHINOIDEA) OF THE GENERA 
MERRIAMASTER AND DENDRASTER FROM 
PURISIMA FORMATION, CALIFORNIA 


By 


J. Wyatt Durham 
Department of Paleontology, University of California, Berkeley, California 94720 


and 
Scott R. Morgan 


Department of Geology, San Francisco State University, San Francisco, California 94132 


ABSTRACT: Merriamaster weaveri n.sp. and Dendraster sullivani n.sp. are described from a sandstone within 
the early Pliocene Tahana Member of the Purisima Formation from a locality near San Gregorio Beach, 
California. Three species (turneri, falorensis, purisimaensis) of Patinopecten (Lituyapecten) occur in se- 
quence in a stratigraphic interval of 45 m at this locality with turneri associated with the echinoids and 
purisimaensis the highest. Fossils in the overlying Pomponio and San Gregorio members suggest a correlation 
of these members with the uppermost type Jacalitos Formation or basal Faunizone F of Adegoke’s 1969 
classification of the Coalinga-Reef Ridge, California sequence. The Tahana Member probably correlates with 


Adegoke’s Faunizone E. 


INTRODUCTION 


Specimens of the sand dollar-type echinoids 
Merriamaster Lambert and Dendraster Agassiz 
are associated in the Tahana Member (Cum- 
mings, Touring and Brabb 1962: 197-200) of the 
Purisima Formation in exposures in the seacliff 
south of San Gregorio Beach, California (Fig. 1). 
This locality is a little over 16 kms south of the 
town of Half Moon Bay. Very probably this is 
the same general locality as that from which 
Scutella perrini Weaver (=Merriamaster perrini 
in current nomenclature) was reported in the 
Santa Cruz Folio (Branner, Newson and Arnold 
1909:6). The same record was subsequently re- 
peated by Kew (1920: 130, as Dendraster perrini) 
and by Grant and Hertlein (1938:68, as Mer- 


riamaster perrini). Weaver (1908:273), in his de- 
scription of S$. perrini, also noted that *‘Other 
specimens closely resembling this form have 
been found at San Gregorio... .”’ J. P. Smith 
(1912:167) listed Astrodapsis perrini Weaver as 
one of ‘the: . characteristic upper Miocene 
species [that] range over into lower Pliocene, 
and become extinct in the Purisima-San Diego 
fauna.’’ Presumably this is Weaver’s Scutella 
perrini, inasmuch as the latter name is used in 
Smith’s checklist on p. 170. Subsequently, 
Smith (1919:145) used the name Dendraster per- 
rini Weaver in his faunal lists for the Purisima. 
Presumably all three names as used by Smith 
refer to the same taxon and were probably based 
on the records by Weaver and by Branner et al. 


[297] 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 11 


Martin (1916:243, and checklist, p. 251) lists 
Scutella perrini Weaver from “* .. . the upper 
standstone [sic] member of the Purisima Forma- 
tion, chiefly from the sea-cliffs south of Half- 
moon Bay.”’ Martin’s collections in the Museum 
of Paleontology at Berkeley (none of the 
California Academy of Sciences collections from 
the Purisima Formation were made by Martin) 
contain no specimens of Merriamaster, so pre- 
sumably the inclusion of M. perrini in his list is 
based on the records by Weaver and by Branner 
et al. We have found no other published record 
of a Merriamaster from the Purisima Formation 
of the type area, and although Cummings et al. 
(1962) recorded Dendraster from the San Gre- 
gorio Member, they did not list it from the 
Tahana Member. However, Carson (1926:55), in 
a largely overlooked paper, lists Dendraster per- 
rini (=Merriamaster perrini) as one of the 
species associated with his Cancellaria palmeri 
and stated that the specimens came from the 
Purisima Formation ‘‘in bluffs above beach east 
of hotel, Capitola, Santa Cruz Co., Calif.’’ This 
material has not been available for examination, 
so it is uncertain whether or not M. perrini was 
correctly identified. The other elements of the 
listed fauna are more suggestive of an upper 
Pliocene age rather than the earlier age inferred 
for the Tahana Member of the type Purisima 
Formation. 

Arnold (1906:27, list) recorded ‘‘Astrodapsis 
n.sp. Merriam’? from the Purisima Formation 
without citing a locality or stratigraphic position 
within the formation. Two years later he (Arnold 
1908:353) listed *‘Astrodapsis, new species, p.” 
from the upper portion of the Purisima Forma- 
tion, stating that ‘*‘These upper beds are typi- 
cally exposed in the sea cliffs in the vicinity of 
Purisima and south to the mouth of Pescadero 
Creek ....”’ This generalized locality descrip- 
tion would include the locality at which the pres- 
ent specimens were collected (as well as the lo- 
cality within the San Gregorio Member of the 
Purisima at which the scutellid echinoids Den- 
draster and Scutellaster occur). Earlier, Haehl 
and Arnold (1904: 18) listed ‘‘Astrodapsis n. sp.”’ 
and (ibid.:24) ‘‘Astrodapsis n. sp. Merriam’’ 
(Merriam never published on this taxon) from 
the Purisima Formation, likewise without spe- 
cific locality citation. Very probably all these ci- 
tations refer to the same species. Further, inas- 
much as Smith (1912:167) assigned Weaver’s 
Scutella perrini to Astrodapsis and Arnold 


acknowledges (1906:8; 1908:345) Smith’s ‘**. . 
assistance in determining the genera and 
zoologic relations of some of the new forms... ,”’ 
it seems possible that all these citations 
(Haehl and Arnold 1904; Arnold 1906, 1908; 
Smith 1912) refer to the same taxon. However, 
because of the lack of descriptions or illustra- 
tions, this suggestion cannot be verified unless 
the collections on which they are based can be 
found. Weathered specimens of Merriamaster 
weaveri N.sp. are suggestive of the genus As- 
trodapsis unless the branching food grooves 
(Fig. 3) on the oral surface are evident, so that an 
assignment to this genus was not unreasonable 
(the genus Merriamaster was not established 
until 1911 and was not generally recognized until 
more than two decades later). 

The specimens on which the present report is 
based were found and collected by the junior 
author of this paper while doing a geologic pro- 
ject under the supervision of Professor Raymond 
Sullivan of San Francisco State University. 
Morgan is responsible for the local stratigraphy 
presented herein, while the senior author is re- 
sponsible for the identification of fossils, de- 
scription of the new species, and stratigraphic 
correlation. 

Some of the fossils from this area have been 
deposited in the Geology Department of the 
California Academy of Sciences (CAS). Most of 
the echinoids, some of the pectinids and a few 
other fossils from the present collection are de- 
posited in the Invertebrate Collections of the 
Museum of Paleontology (UCMP) of the Uni- 
versity of California (Berkeley). The Museum 
of Paleontology collections contain numerous 
other fossils from the same general area as the 
echinoids, but they are of less certain relative 
stratigraphic positions. 


STRATIGRAPHY 


The bed with the echinoids (the echinoids and 
some associated fossils are catalogued in the 
Museum of Paleontology under locality number 
D 3399) is exposed in the seacliff about 760 m 
south of the parking lot at San Gregorio Beach. 
It is a medium fine-grained, grey-green (un- 
weathered color) sandstone bed about 30 cm 
thick, with the sand dollars in irregularly distrib- 
uted concentrations. Some of the echinoids were 
crushed or broken before burial, indicating 
some reworking before final entombment. The 
echinoid-bearing bed is exposed in the axial area 


DURHAM & MORGAN: NEW SAND DOLLARS 


San Francisco o 


San Gregorio Beach 


Pomponio Beach 


FIGURE I. 


of a gentle east-west trending anticline and at a 
maximum attains a height of about 3 m above the 
base of the cliff. It gradually disappears below 
the beach level to both the north and south. The 
local rock sequence (Fig. 2) is referable to the 
upper part of the Tahana Member of the 
Purisima Formation (see Cummings et al. 
1962:197-200, pl. 20). The part of the Purisima 
Formation exposed (Fig. 1) between the axis of 
the anticline and San Gregorio Creek to the 
north is about 76 m thick and is composed of 
massive marine sandstones with some interbed- 


299 


meters 


Map of San Gregorio Beach area, showing location of fossil collecting localities (‘‘D”’ numbers). 


ded siltstone and a rhyolitic tuff. A horizontal 
Quaternary non-marine terrace conglomerate up 
to 7 m thick unconformably overlies the older 
sequence. Fossil horizons generally correspond 
to concretionary beds or to the more calcareous 
strata. Two other fossiliferous beds (localities 
UCMP D 3400 and D 3401) occur below the 
echinoid-bearing stratum, with the collections 
being made on both sides of the anticlinal axis. 
Two other significant collections (UCMP locs. D 
6345 and D 6346) from higher in the sequence 
were made from the strata exposed to the south 


50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 11 


50 
+D6346 
” 
254 = 
@ 
fe 
+D6345 
10 
+D3399 0 


+D3400 
+D4301 


Scott R. Morgan 
December 27, 1976 


Columnar section of the Tahana Member of the 
Purisima Formation and overlying Quaternary terrace de- 
posits as exposed in seacliffs between Pomponio and San 
Gregorio creeks, San Mateo County, California. ‘‘D’’ prefix 
numbers represent fossil collections in the Museum of Paleon- 


FIGURE 2. 


of the axis. The same beds do not appear to be 
fossiliferous on the north flank of the anticline. 

The sand dollars at locality D 3399 are as- 
sociated with other fossils, including Swifto- 
pecten swifti parmeleei (Dall), Patinopecten 
(Lituyapecten) turneri (Arnold) (hypotype 
UCMP 14457), Spisula albaria (Conrad), Yoldia 
scissurata subsp. strigata (Dall), Nassarius 
(Caesia) grammatus (Dall), and Clinocardium 
nuttallii (Conrad)(?). 

In the bed immediately below (UCMP D 3400) 
the echinoid-bearing stratum, the barnacle, 
Balanus (Tamiosoma) gregarius (Conrad), is as- 
sociated with Swiftopecten swiftii parmeleei. In 
a bed (loc. D 3401) about 3 to 3.5 m below the 
echinoids, there are abundant pectens that have 
been identified as Patinopecten healeyi (Arnold) 
(see Arnold 1906:104, pl. 37, fig. 2 only; and 
Cummings et al.: photo 15, fig. 4). All pectens 
from this bed are like those illustrated by Arnold 
and by Cummings et al. and do not agree well 
with the most common variants of P. healeyi 
(see Hertlein and Grant 1972: pl. 21, figs. 1, 6, 7) 
from the San Diego Formation. About 16 m 
stratigraphically above the echinoid bed (UCMP 
14458, loc. D 6345) a specimen of Patinopecten 
(Lituyapecten) falorensis MacNeil was found. 
P. (L.) falorensis has previously been reported 
from this area by MacNeil (1961:234—235). The 
locality description for his specimens is not pre- 
cise stratigraphically but it suggests a position 
similar to that of the present specimen. About 41 
m stratigraphically above the echinoid-bearing 
bed, specimens of P. (L.) purisimaensis (Arnold) 
(UCMP 14459, 14260, loc. D 6346) are present. 
Thus the local evidence suggests that P. (L.) 
furneri, P: (L:) falorensis, ‘and P(e) 
purisimaensis form a sequence, occurring in that 
order from oldest to youngest. 

Elsewhere Patinopecten (Lituyapecten) tur- 
neri occurs in the Merced Formation in the Es- 
tero San Antonio area (type area) near Dillon 
Beach, California (see Peck 1960: pl. 21, figs. 
15-16). Peck (1960: table 2, pl. 21, fig. 14 [labeled 
as left valve but actually right valve]) also re- 
corded P. (L.) purisimaensis from the Ohlson 
Ranch Formation near the San Andreas Fault 
in northwestern Sonoma County, California. 
Reexamination of the illustrated specimen and 


— 


tology (UCMP), University of California (Berkeley), and their 
stratigraphic positions. 


DURHAM & MORGAN: NEW SAND DOLLARS 


additional fragmentary specimens indicates 
that they should be referred to P. (L.) falorensis . 
To date, typical P. (L.) purisimaensis has been 
validly recorded from formations other than the 
Purisima only by Glen (1959: 168, pl. 15, figs. 2, 
3) from the *‘Merced Formation’’ of Pillar Point 
(about 22 km northwest of the Merriamaster lo- 
cality). Elsewhere P. (L.) falorensis occurs in 
the Falor Formation (type-specimens) and the 
Rio Dell Formation of Humboldt County, 
California (MacNeil 1961:235). 


AGE 

Cummings et al. (1962:211) considered the 
Tahana Member of the Purisima Formation to be 
near the base of the Pliocene Series in the sense 
of the traditional usage of Pliocene in California. 
The formation unconformably overlies beds as- 
signed to the Monterey Formation and which 
Cummings et al., (1962:197) suggested were 
younger than Luisian and possibly correlative 
with the Delmontian portion of the type Mon- 
terey Formation. 

The Tahana Member is overlain by the Pom- 
ponio Member, and it in turn by the San Gre- 
gorio Member of the Purisima Formation. Fos- 
sils reported by Cummings et al. (1962:202—208) 
from these two members include Patinopecten 
lohri (Hertlein), Anadara trilineata (Conrad), 
and Dendraster gibbsii (Remond), suggesting a 
correlation with the upper Jacalitos-Etchegoin 
succession of the Coalinga-Kettleman Hills area 
of the San Joaquin Valley (upper part of 
Faunizone E and Faunizone F of Adegoke 
1969:76-80). The Scutellaster oregonensis of 
Cummings et al. (1961: photo 19, fig. 5) is not 
now referred to that species but is closely similar 
to a new species (originally identified as S. 
oregonensis quaylei in Durham and Wolfe 1958) 
from the uppermost beds of the type Jacalitos 
Formation near Coalinga. In the Jacalitos oc- 
currence, Scutellaster is associated with Den- 
draster gibbsii, while Patinopecten lohri oc- 
curs in nearby strata. Thus the Scutellaster 
suggests that the Pomponio-San Gregorio mem- 
bers are correlative with the uppermost part of 
the type Jacalitos Formation or zonule 10 of 
Faunizone F of Adegoke (1969:78-79) of the 
Coalinga-Reef Ridge area. 

The three species of Patinopecten (Lituyapec- 
ten) from the Tahana Member suggest that it can 
be correlated with parts of the Falor Formation, 
the Ohlson Ranch Formation and the Merced 


301 


Formation near Dillon Beach and Bodega Bay. 
The stratigraphic position of the Tahana 
Member below the Pomponio Member suggests 
that it is older than the uppermost type Jacalitos 
Formation or Adegoke’s Faunizone F of the 
Coalinga-Reef Ridge area, and that it is probably 
to be correlated with the older part of the type 
Jacalitos Formation or Faunizone E of 
Adegoke’s terminology. In traditional West 
Coast megafaunal terminology, the age of the 
Tahana Member is early Pliocene, but it should 
be recognized that correlations based on 
planktonic foraminifera may show that it is of 
late Miocene age in terms of the European type 
sections. 


SYSTEMATIC DESCRIPTIONS 
Genus Merriamaster Lambert, 1911 


Merriamaster weaveri n.sp. 
(Figure 3) 
2Scutella perrini Weaver in Branner, Newson and Arnold 


1909:6; Weaver 1908:273(in part); non Weaver 1908:273(in 
part), pl. 22, fig. 2; nec auctores. 


Diagnosis.—A large Merriamaster with less 
eccentric apical system, slightly longer and less 
flared posterior petals, and sparser and finer 
tuberculation than in M. perrini; tuberculation 
not as fine as in M. arnoldi. 

Description.—Adult test of large size for 
genus, margin moderately inflated; position of 
apical system variable, slightly eccentric; 
maximum height of test anterior to apical system; 
petals elongate, distally with nearly parallel 
sides, sometimes raised adjacent to apical sys- 
tem; length of anterior petal about 80 percent of 
corresponding radius; periproct just submargi- 
nal; peristome slightly posterior; numerous mod- 
erately large tubercles within petals, tubercles in 
interambulacra moderately fine; internal concen- 
tric test supports 2 to 4 (rarely) as in type-species. 

Dimensions .—Holotype (UCMP 14259) length 
38.8 mm, width 37.4 mm, thickness 11.0 mm; 
paratype (UCMP 14443) length 47.8 mm, width 
+40 mm, thickness about 9-10 mm; paratype 
(UCMP 14445), length +51 mm, width +48.8 
mm, thickness +10.2 mm; paratype (UCMP 
14446), length +37 mm, width 33.5 mm, height 
7.3 mm; paratype (UCMP 14449) crushed, poorly 
preserved, major observed diameter +62 mm. 

Types .—All from UCMP loc. D 3399; holotype 
UCMP 14259, paratypes UCMP 14443 to 14447, 
14449; paratypes CAS 58175 and 58176. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 11 


SION Bee A ie 


ae 


FIGURE 3. Merriamaster weaveri n.sp. A-D, F, approximately x1; E, x 1.4; G, x3.4. (A) Paratype, UCMP 14443, part of 
another specimen on right side. (B) Paratype, UCMP 14445, immersed in water. (C) Paratype, CAS 58176. (D) Paratype, UCMP 
14446, immersed in xylene. (E) Holotype, UCMP 14259, apical surface, note large tubercles in ambulacra. (F) Paratype, UCMP 
14447. Traces of tuberculation in ambulacrum V. (G) Part of oral surface of same paratype as fig. D. Ambulacrum I in left 


DURHAM & MORGAN: NEW SAND DOLLARS 


Occurrence.—UCMP loc. D 3399 (only), early 
Pliocene. 

Discussion.—There are 10 reasonably well- 
preserved specimens, but some of them are 
crushed or incomplete. There are also numerous 
poorly preserved individuals. The species is 
named after C. E. Weaver who described the 
type-species (Scutella perrini) of the genus. It 
differs from M. perrini by the less eccentric apical 
system and the straighter, longer, and more open 
posterior petals. The tuberculation is coarser 
than that of M. arnoldi (Twitchell) which is nearly 
always associated with M. perrini. It differs from 
the species represented by the specimen from the 
Sargent Oil Field figured by Kew (1920:pl. 28, 
figs. 2a, 2c) as Dendraster arnoldi (now Mer- 
riamaster arnoldi) by the less eccentric apical 
system and sparser tuberculation within the 
aboral interambulacral areas of the new species. 
Merriamaster cf. M. perrini (Weaver) (Woodring 
and Bramlette 1950:pl. 10, figs. 11-14) from the 
Foxen Mudstone of the Santa Maria District dif- 
fers by the sparser tubercles on the oral surface 
and its nearly central apical system. 

Merriamaster weaveri differs from the as- 
sociated Dendraster sullivani by its less eccentric 
apical system, by its higher (radial dimension) 
interambulacral plates on the apical surface, and 
by its thicker margin. 

Merriamaster weaveri is of older age than M. 
perrini and M. arnoldi from the late Pliocene of 
the San Joaquin Valley area, and presumably is 
older than M. pacificus (Kew, 1920) from the San 
Diego area, as well as M. israelskyi (Jordan and 
Hertlein, 1926) and M. kewi (Jordan and Hertlein, 
1926) of the Pliocene of Cedros Island, Baja 
California, all of which have usually been as- 
signed a ‘‘middle’’ Pliocene age. It may represent 
the ancestral stock of the genus. 


Dendraster sullivani n.sp. 
(Figures 4, 5.) 


Diagnosis .—An intermediate-sized Dendras- 
ter with thin margins in large adults; bivium angle 
about 90°; petals nearly wide open, poriferous 
zones wide; periproct submarginal. Large tuber- 
cles within petals. 

Description.—Adult test of medium size, with 
thin margins; apical system moderately eccen- 


— 


303 


tric, distant about 38-40% of length from poste- 
rior margin; maximum thickness just anterior to 
apical system; posterior petals wide, about % 
length of corresponding radius, only very slightly 
arcuate with very slight tendency to close, pore- 
zones tapering slightly distally; anterior petal 
narrow, abruptly constricted about 4 distance 
from distal end, then continuing nearly parallel 
sided; some demi-ambulacral plates in con- 
stricted area; anterior paired petals nearly paral- 
lel sided in distal 24, then abruptly constricted to 
end; oral surface slightly concave, peristome just 
slightly posterior; tubercle pattern on oral surface 
not completely discernible but moderately nu- 
merous large tubercles in central % of surface; 
tuberculation on aboral surface mostly destroyed 
but some remnants present, tubercles fewer and 
more elevated than on D. excentricus and D. 
gibbsii; a few large tubercles inside poriferous 
area of petals as in D. vizcainoensis; periproct 
submarginal, in an admarginal position between 
second pair of post-basicoronal interambulacral 
plates. 

Dimensions .—Holotype, length +51 mm (pos- 
sibly 1-2 mm missing), width +55 mm (1-2 mm 
missing); height, + 8 mm; paratype 14455, very 
incomplete, probably about 65 mm wide. 

Types.—All from UCMP loc. D 3399. Holo- 
type, UCMP 14450, paratypes UCMP 14448, 
14451 to 14456; paratypes CAS 58177 to 58179. 

Occurrence.—UCMP loc. D 3399 only, early 
Pliocene. 

Discussion.—Named after Professor Ray- 
mond Sullivan. This species differs from D. 
excentricus (Agassiz) and D. gibbsii (Remond) by 
its coarser tuberculation, larger tubercles within 
the petals, and thinner margin. It differs from D. 
elsmerensis Durham of similar age, which 
likewise has a thin margin, by its submarginal 
instead of marginal periproct, its smaller bivial 
angle, and by the presence of large tubercles 
within its petals (see paratypes CAS 58177 and 
UCMP 14453). The abrupt constriction of the 
anterior petal seems to be unique. The large 
tubercles suggest that it belongs to the group of 
Dendraster vizcainoensis Grant and Hertlein 
(1938:90, pl. 8, figs. 1-3). 

The termination of the anterior petal, with the 
abrupt constriction and demi-plates, is unusual 


quadrant, note fine ‘“‘tuberculation”’ of food groove branching about midway to ambitus. Irregular radial fracture near middle of 
quadrant is perradial suture (bisecting food groove). Peristome admarginal in lower left. 


4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 11 


Ficure 4. Dendraster sullivani n.sp. A, B, D, G, approximately x1; C, x5; E, F, approximately x3.5. (A) Holotype, 
UCMP 14450, immersed in xylene. (B) Paratype, UCMP 14451. (C) Paratype, UCMP 14453, note remains of large tubercles 
within petal, and small tubercles outside in interambulacrum. (D) Paratype, UCMP 14452. (E) Paratype, CAS 58177, note badly 
eroded large tubercles in center of petal and a few small tubercles preserved in lower right. (F) Same specimen as D, note poorly 


DURHAM & MORGAN: NEW SAND DOLLARS 


L 79 mm 1 


Ficures 5. Dendraster sullivani n.sp., paratype UCMP 
14454. Distribution of pores and plates around tip of anterior 
petal. Note demi-plates around tip of petal. 


for this genus, but it is clearly displayed on the 
holotype and paratype 14454, the only specimens 
on which this area is well preserved. 

The exterior surface of the plates has been 
eroded or removed on most specimens, with ex- 
ternal tuberculation being preserved in only afew 
patches. Growth lines are well shown on several 
specimens, a condition which is rare in species 
of Dendraster. Presumably these well-defined 
growth lines indicate a marked periodicity of the 
conditions that caused their formation. On 
intermediate-sized individuals (diameter about 40 
mm), the margin is not as thin as on larger speci- 
mens. 


LITERATURE CITED 


ADEGOKE, O. S. 1969. Stratigraphy and paleontology of the 
marine Neogene formations of the Coalinga Region, 
California. Univ. Calif. Publ. Geol. Sci. 80: 1-241, pls. 1-13. 

ARNOLD, RALPH. 1906. The Tertiary and Quaternary pec- 
tens of California. U.S. Geol. Surv. Prof. Pap. 47:1-264, 
pls. 1-53. 

1908. Description of new Cretaceous and Tertiary 

fossils from the Santa Cruz Mountains, California. Proc. 

U.S. Natl. Mus. 34:345-390, pls. 31-37. 


— 


305 


BRANNER, J. C., J. F. NEWSON AND RALPH ARNOLD. 1909. 
Santa Cruz folio. California. Geol. Atlas United States, no. 
163. U.S. Geol. Surv. 11 pp., maps. 

CARSON, CARLTON M. 1926. New molluscan species from 
the California Pliocene. Bull. South. Calif. Acad. Sci. 
25:49-62, pls. 1-4. 

CUMMINGS, J. C., R. M. TOURING AND E. E. BRABB. 1962. 
Geology of the northern Santa Cruz Mountains, California. 
Calif. Div. Mines Geol. Bull. 181:179-220, pls. 20-24. 

DuRHAM, J. W., AND J. A. WOLFE. 1958. Joint occurrence 
of Dendraster and Scutellaster [Anorthoscutum]. Bull. 
Geol. Soc. Am. 69: 1682-1683. 

GLEN, WILLIAM. 1959. Pliocene and lower Pleistocene of 
the western part of the San Francisco peninsula. Univ. 
Calif. Publ. Geol. Sci. 36:147—198, pls. 15-17. 

GRANT, U. S., IV, AND L. G. HERTLEIN. 1938. The West 
American Cenozoic Echinoidea. Publ. Univ. Calif. Los 
Angeles Math. Phys. Sci. 2:1—225, pls. 1-30. 

HaeEHL, H. L., AND RALPH ARNOLD. 1904. The Miocene 
Diabase of the Santa Cruz Mountains in San Mateo County, 
California. Proc. Am. Philos. Soc. 43(175): 15-53. 

HERTLEIN, L. G., AND U. S. GRANT, IV. 1972. The geology 
and paleontology of the marine Pliocene of San Diego, 
California (Paleontology: Pelecypoda). San Diego Soc. Nat. 
Hist. Mem. 2 (pt. 2B):143-409, pls. 27-57. 

JORDAN, E. K., AND L. G. HERTLEIN. 1926. Contribution to 
the geology and paleontology of the Tertiary of Cedros Is- 
land and adjacent parts of Lower California. Proc. Calif. 
Acad. Sci., Ser. 4, 15:409-464, pls. 27-34. 

Kew, W. S. W. 1920. Cretaceous and Cenozoic Echinoidea 
of the Pacific coast of North America. Univ. Calif. Publ. 
Bull. Dep. Geol. 12:23-236, pls. 3-42. 

MACNEIL, F. S. 1961. Lituyapecten (new subgenus of 
Patinopecten) from Alaska and California. U.S. Geol. Surv. 
Prof. Pap. 354:225—239, pls. 3546. 

MarTIN, BRUCE. 1916. The Pliocene of middle and northern 
California. Univ. Calif. Publ. Bull. Dep. Geol. 9:215-259. 
Peck, J. H., Jk. 1960. Paleontology and correlation of the 
Ohlson Ranch Formation. Univ. Calif. Publ. Geol. Sci. 

36:235—242, pl. 21. 

SMITH, J. P. 1912. Geologic range of Miocene invertebrate 
fossils of California. Proc. Calif. Acad. Sci., Ser. 4, 3:161- 
183. 


1919. Climatic relations of the Tertiary and Quarter- 
nary faunas of the California region. Proc. Calif. Acad. Sci., 
Ser. 4, 9:123-173. 

WEAVER, C. E. 1908. New echinoids from the Tertiary of 
California. Univ. Calif. Publ. Bull. Dep. Geol. 5:271-274, 
pls. 21-22. 

WoOoDRING, W. P., AND M. N. BRAMLETTE. 1950. Geology 
and paleontology of the Santa Maria District, California. 
U.S. Geol. Surv. Prof. Pap. 222:1-185, pls. 1-23. 


preserved tubercles in upper central petal (Lovenian no. I). (G) Paratype, CAS 58178, weathered oral surface, interambulacrum 
5, center of lower margin. Note poorly preserved large tubercles towards peristome. 


Mer ig th) i ie i 
—. f wecrraia fe . oon 
the custom. Pacilix (nas ‘y Roy 
ides (UTA) Geuli os 1! ¥ 7 : | ae aia 
aoe Miihigesere i's me ised rahi ans 
a 0c Tage 


Palliat: 1 y ws : 
i" » Cems? Gets MW ny fh 


ee tee ae va ee eee 
ao uta wh baaged > at a 


mat ligt . 
r sch ty“ Teniehogn; t Amy 


fore oe 


GOs inal 
i 
-— 


PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 12, pp. 307-337; 20 figs. 


February 8, 1978 


EASTERN PACIFIC MACROURIDS OF THE GENUS 
COELORINCAHUS GIORNA (PISCES: GADIFORMES), WITH 
DESCRIPTION OF A NEW SPECIES FROM CHILE 


By 


Tomio Iwamoto 
California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 


ABSTRACT: Six eastern Pacific representatives of the speciose and widely distributed grenadier genus Coelorin- 
chus are known. One of these, C. aconcagua, is herein described as new; it is a species of relatively shallow 
depths (175—428 m) and restricted distribution (off Chile between approximately latitudes 30° and 42° S). Three 
other species also have relatively restricted distributions: C. scaphopsis (southern California and northern Gulf 
of California); C. canus (northern Peru to Costa Rica); and C. chilensis (northern Chile to Peru). C. innotabilis 
and C. fasciatus are widely distributed in cold-water regions of the southern hemisphere; one Chilean specimen 
of the former represents a first record from the eastern Pacific. Five populations of C. fasciatus show slight 
morphological differences from each other; a clinal pattern is seen in several features from South America to 
South Africa to South-West Africa. C. patagoniae Gilbert and Thompson, 1916, is synonymized with C. 
fasciatus. C. canus and C. scaphopsis appear most closely related to western Atlantic species; C. fasciatus is 
widespread in the subantarctic and has close relatives in Australian waters; C. innotabilis appears most closely 
related to members of the subgenus Paramacrurus from the Philippines, East Indies, Japan, and Hawaii; the 
relationships of C. chilensis and C. aconcagua are obscure. 


INTRODUCTION 


This paper is the fourth in a series dealing with 
the systematics of macrourid fishes, or gren- 
adiers, of the eastern Pacific Ocean. The first 
(Iwamoto and Stein 1974) dealt with 10 species of 
grenadiers of the northeastern Pacific north of 
San Francisco, California. The other two 
(Chirichigno and Iwamoto 1977; and Hubbs and 
Iwamoto 1977) reported new species from the 
eastern Pacific. The purpose of this paper is to 
treat systematically the six eastern Pacific repre- 
sentatives of the speciose genus Coelorinchus , 
discussing their relationships and distribution 
and describing one as new and another as a first 
record from the region. 


MATERIAL AND METHODS 


Papers of this series are based primarily on 
studies of the extensive material collected within 
the past two decades by vessels of several United 
States institutions. Collections made by oceano- 
graphic research vessels of Oregon State Univer- 
sity formed the bulk of the material reported on 
by Iwamoto and Stein (1974) (most of that mate- 
rial is now housed at the California Academy of 
Sciences, CAS). The extensive midwater collec- 
tions made by vessels of the Scripps Institution of 
Oceanography (SIO) and the University of 
Southern California (latter collections in Los 
Angeles County Museum of Natural History, 
LACM) formed the basis of the Hubbs and 


[307] 


08 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


Iwamoto (1977) article describing new bathy- 
pelagic grenadiers. 

For the present article and for subsequent arti- 
cles dealing primarily with species of the warm- 
water and southern-hemisphere regions of the 
eastern Pacific, the macrourids captured by the 
Smithsonian Institution’s ANTON BRUUN are 
preeminent among recent collections. Represen- 
tatives of at least 19 of the approximately 27 
species known from the Pacific coast of South 
America were taken by that vessel in 1966 during 
cruises involved with the Southeastern Pacific 
Biological Oceanography Program. Supplement- 
ing the ANTON BRUUN collections are the sub- 
stantial ones made by Stanford University’s TE 
VEGA (collections housed in CAS), the Univer- 
sity of Southern California’s VELERO IV, and 
various vessels of the Scripps Institution of 
Oceanography. 

Of all the eastern Pacific grenadier collections, 
those made by the United States Fish Commis- 
sion steamer ALBATROSS around the turn of 
the century must stand foremost. Most of the 
ALBATROSS grenadiers from southern- 
hemisphere and warm-water regions of the east- 
ern Pacific have already been treated by Garman 
(1899), Gilbert (1890; 1892), and Gilbert and 
Thompson (in Thompson 1916). Important col- 
lections made by that vessel in 1888 and 1904 in 
waters off the Galapagos, mainland Equador, and 
Panama have, however, never been reported on, 
although Dr. Carl. L. Hubbs (SIO) examined the 
material while a student at Stanford University 
and applied names (some new) to the specimens 
before returning most (some were retained at 
Stanford) to the National Museum of Natural 
History (USNM). Because of other commit- 
ments, he was never able to get back to complete 
a publication on them (Carl L. Hubbs, personal 
communication). Most of the ALBATROSS col- 
lections from the eastern Pacific are housed in the 
USNM, Museum of Comparative Zoology at 
Harvard University (MCZ), and CAS (CAS-SU 
refers to specimens formerly housed at the Natu- 
ral History Museum of Stanford University). 

Specimens examined from institutions other 
than those listed above and used for the present 
paper include: American Museum of Natural His- 
tory, New York, N.Y. (AMNH), Gulf Coast Re- 
search Laboratory Museum, Ocean Springs, 
Mississippi (GCRL), Instituto del Mar, Callao, 
Peru (IMARPE), Musée Royal de I’ Afrique Cen- 
trale, Tervuren, Belgium (MRAC), Museo Na- 


cional de Historia Natural, Santiago, Chile 
(MNHN, Santiago), and South African Museum, 
Cape Town, South Africa (SAM). 

Methods for making counts and measurements 
follow procedures outlined in a previous work 
(Iwamoto, 1970) and generally follow the meth- 
odology of Hubbs and Lagler (1958). Figure 1 
illustrates how certain measurements were 
taken. Because of the importance of certain 
counts as diagnostic characters, methods for tak- 
ing them are here described in detail: 


First dorsal fin—counts include the two spinous 
anteriormost rays (the first of which is small, 
closely appressed to the long second ray, and 
scarcely discernible without dissection). They 
are designated in counts given in the text by 
Roman numerals but are not differentiated in 
the tables. The last two rays of the fin are 
usually unbranched, and each is included in the 
fin-ray count. 

Pectoral fin—has a small splintlike uppermost 
ray which Is designated by the letter i, in lower 
case, in counts given in the text but is not 
differentiated in the tables. The lowermost rays 
of the fin are very small and require substantial 
magnification and often teasing of the folds of 
the skin at the fin base to count them accu- 
rately. 

Lateral-line scales over distance equal to predor- 
sal length—the number of lateral-line scales, 
counted from the anterior origin of the lateral 
line, over a distance equal to that of the predor- 
sal length. 


Scales below 1D.—Scale rows above the lateral 
line, counted obliquely down and back from 
the origin of the first dorsal fin to the lateral 
line, but not including the lateral-line scale. 
The small size of the scales at the dorsal origin 
and the irregularity of rows there make accu- 
rate counts difficult. Designation of half scales 
in the counts indicates that the scale nearest the 
median line is much smaller than the adjacent 
lateral (lower) scale. 

Scales below mid-1D., and below 2D.—takenina 
similar manner as for scales below 1D. except, 
in the first case, the count is taken below the 
midbase of the first dorsal fin while, in the 
second case, it is taken below the origin of the 
second dorsal fin. 


Gill-rakers—all rakers on the inner (medial) side 
of the arch, including rudimentary ones, on the 
upper and lower limbs are counted. In the text, 


IWAMOTO: EASTERN PACIFIC MACROURIDS 309 


Predorsal 


Pectoral 


(i 
L-- 


Preoral 


Pelvic <__ 


\ Preoral 5 
; ">> Snout 


FicureE |. Methods for taking certain measurements in specimens of Coelorinchus: a) lateral view: b) ventral view; c) dorsal 
view. Abbreviations: Barbel—length of barbel; 1D.—height of first dorsal fin; 1D.-2D.—length of space between first and second 
dorsal fins; HL—head length; Intorb.—least width of interorbital space; Istm—A.—distance from isthmus to anal origin; Nasal— 
least (internasal) width between nasal ridges (taken from outer edge of ridges); pectoral—length of pectoral fin; Pelvic—length of 
pelvic fin; 2P.-A.—distance between base of outer pelvic ray and anal origin; Predorsal—predorsal length; Preoral—preoral 
length of snout; Postorb.—least postorbital length of head; Orbit—greatest orbit diameter; Orb—Preop.—distance between 
posterior margin of orbit and posterior margin of preopercle; Snout—length of snout; Suborb.—least width of suborbital region. 


310 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


FIGURE 2. 
head of Coelorinchus showing positions of head ridges. Ab- 
breviations: front.—frontal ridge; med. rost.—median rostal 


Diagrammatic dorsal (a) and lateral (b) views of 


ridge; par.—parietal ridge; postorb.—postorbital ridge; 
suborb.—suborbital ridge; | supn.—supranarial _ ridge; 
supoc.—supraoccipital ridge; suporb.—supraorbital ridge. 


counts of the upper limb are usually separated 
from those of the lower limb by a plus (+) sign. 
In some cases the uppermost and lowermost 
rakers are minute and require substantial mag- 
nification and teasing of the surrounding tissue 
to expose them. 


Head ridges in species of Coelorinchus are par- 
ticularly prominent compared with those found in 
other genera of macrourids. The ridges are 
sharply defined by rows of stout, spiny, deeply 
embedded, nonimbricate, scutelike scales. Of- 
ten, the spinules on the ridge scales.are imbricate, 
aligned in longitudinal or divergent rows, and 
form sharp, serrated keels or crests. Terminology 
for the head ridges generally follows Okamura 
(1970a:6—7, text-fig. 2) with some modifications. 
Names and diagrams of the positions of the ridges 
as I use them for Coelorinchus are shown in Fig- 
ure 2. I use the term suborbital to collectively 
refer to Okamura’s three separately defined 
ridges—the lateral nasal, infraorbital, and the 
longitudinal axis of the preopercular ridge. These 


three ridges in Coelorinchus form essentially one 
continuous ridge, and I have found it more con- 
venient to refer to them here as a single unit. 

In the ‘‘Materials Examined”’ sections, collec- 
tions are listed according to geographic locality 
and latitude, the northernmost captures for each 
political entity listed first. Museum catalog num- 
bers are followed (in parentheses) by the number 
of specimens and their range of head length 
and/or total length (in mm). A plus sign after 
a number indicates the specimen had a termi- 
nal portion of the tail missing (whether a 
‘*pseudocaudal”’ is developed or not). The geo- 
graphical coordinates, the capture depths, and 
other pertinent capture data follow. 


Genus Coelorinchus Giorna 


Coelorinchus Giorna, 1809:177—180 (type-species Coelorin- 
chus La-Ville Risso, 1810, by monotypy). 


D1AGNosis.—Macrourine grenadiers with six 
branchiostegal rays. Abdominal light organ usu- 
ally well developed, often with one or two large 
external dermal windows anterior to, and often 
very remote from, anus. Anus usually im- 
mediately in advance of anal fin or much closer to 
anal-fin origin than to pelvic bases. Second spi- 
nous ray of first dorsal fin usually smooth, rarely 
with a few small denticles near tip. Suborbital 
ridge strongly developed, usually very angular in 
cross section. Head ridges strong, reinforced 
with stout scutelike scales. Snout moderately to 
greatly projecting. Orbits large, elliptical, usually 
less than 1.5 into postorbital length of head. Body 
relatively shallow, belly region broad and long. 
Pelvic fins widely separated, consistently with 7 
rays. Interopercle usually completely hidden be- 
hind preopercle. 

REMARKS.—Okamura (1970a:148) gave a de- 
tailed description of the genus using additional 
generic characters not in the above diagnosis. A 
broader survey of the genus, however, will likely 
necessitate modification of some of his diagnostic 
characters. 

See Marshall and Iwamoto (in Marshall 
1973:538) for a generic synonymy. Okamura 
(1970a:143; 1970b) has recognized Abyssicola 
Goode and Bean, 1896, with its only species A. 
macrochir (Ginther, 1877), as distinct from 
Coelorinchus based on the following features: 
large mouth; weak infraorbital ridge, strongly 
curved anteriorly; suborbital region not horizon- 
tal; head compressed; teeth in series; and anus 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


remote from anal fin. Stated thusly, these fea- 
tures do not truly reflect the very different 
physiognomy of A. macrochir , as compared with 
that of most species of Coelorinchus. In particu- 
lar, the structure of the suborbital ridge, the rela- 
tively large mouth, and the dentition are quite in 
contrast with the same features in species of 
Coelorinchus. But the relationships of this 
monotypic genus are obviously with Coelorin- 
chus (to which Okamura agrees), and they share 
so many features in common that one must won- 
der if subgeneric treatment of Abyssicola, follow- 
ing Gilbert and Hubbs (1920:425), might not be 
preferable. 

Members of the genus Coelorinchus form a 
tightly knit group within the large and diverse 
subfamily Macrourinae. As far as known, all 
members of the genus are closely associated with 
the bottom as adults, with forays into the overly- 
ing water column probably infrequent (C. asper- 
cephalus Waite, 1911, with its terete body and 
rounded snout, may be an exception). The gen- 
eral external morphology in most species of 
Coelorinchus suggests a bottom dweller. The 
mouth is inferior, often very small and U-shaped, 
the ventral aspects of the head and body are 
somewhat flattened, the gill openings are usually 
restricted ventrally, the suborbital ridge is rein- 
forced by heavy scutelike scales, and the snout is 
usually pointed and spade-shaped. 

An abdominal light organ is present in all 
species of Coelorinchus. It has been studied in 
detail by several workers but most notably by 
Haneda (1951), Hickling (1931), and Okamura 
(1970a, 1970b). The saclike organ, which houses 
symbiotic luminescent bacteria, is situated within 
the body wall in front of the anus. The lumen of 
the organ is connected to the rectum via a small 
duct through which the luminescent bacteria can 
presumably be ejected to the exterior. Hickling 
(1931) believed that the organ in C. coelorhincus 
is fully functional in the young but becomes es- 
sentially nonfunctional in older individuals. The 
organ has been of taxonomic use in the family 
Macrouridae since Gilbert and Hubbs (1916) first 
recognized it as being of possible luminescent 
function. In Coelorinchus it is variously devel- 
oped, from a small, short structure abutting the 
anterior wall of the anus, to an elongated double- 
sacced structure with one sac located on the chest 
between the pelvic fins and connected by a long 
duct toa second slightly smaller sac in front of the 
anus. Separation of the five nominal subgenera of 


311 


gas gland 


rete 


«#— drumming —» 
muscles 


Ficure 3. Diagrammatic illustrations of gasbladder of 
six species of Coelorinchus showing general shape, position of 
oval window, shape and structure of rete-gas gland complex 
and, in two species, position of drumming muscles: (A) C. 
fasciatus, 2, 52mm HL, CAS-SU 23006; (B) C. occa, 2, 101 
mm HL, CAS 14529; (C) C. chilensis, 6, 69 mm HL, CAS 
uncat.; (D) C. aconcagua, ?, 583 mm HL, CAS 36801; (E) C. 
canus, 2, 34 mm HL, CAS 35914; (F) C. scaphopsis, 2 , 46 
mm HL, CAS-SU 179. Scales represent 5 mm. 


Coelorinchus is based primarily upon the relative 
development of the organ. The sequence from 
lowest to highest development is seen in the sub- 
generic series: Oxygadus, Oxymacrurus, Para- 
macrurus, Coelorinchus, Quincuncia. 


312 


The gas bladder of Coelorinchus is large and 
usually oval, but in a few species it is partially or 
completely divided into anterior and posterior 
chambers with the anterior chamber bilobed. C. 
fasciatus (Gunther, 1878) and its closest relatives 
exhibit this latter condition. Species of Coelorin- 
chus generally, but not invariably, have four large 
gas glands, each attached to a short broad rete. In 
C. fasciatus the rete-gas gland complex is usually 
divided into two pairs, one pair for each lobe of 
the anterior chamber. In nine C. canus (Garman, 
1899) specimens I examined, the number of rete- 
gas gland combinations varied from 6-11; gener- 
ally the higher the number, the smaller the size of 
each combination. Gas bladders of five eastern 
Pacific species and one Atlantic species of 
Coelorinchus are diagrammed in Figure 3. 

Sexual dimorphism has been noted in certain 
species of Coelorinchus. Gilbert and Hubbs 
(1920:371) reported that the first dorsal, pectoral, 
and ventral fins of C. velifer Gilbert and Hubbs, 
1920, and its allies are longer in adult males than 
in females or young males. Marshall (1965:313) 
found well-developed drumming muscles on the 
gas bladder in males of various species of 
Coelorinchus (and other macrourid genera) but 
not in the females. 

The genus Coelorinchus is the most speciose of 
the approximately 25 genera of the Macrouridae. 
The taxon reaches its greatest diversity in the 
Philippines, where Gilbert and Hubbs (1920:424) 
reported 23 species, or 45 percent of the (then) 
known species of the genus. Off Japan the genus 
shows a lesser, but still high, diversity with 16 
species (Okamura 1970a). Other major faunal 
areas have fewer species: Australia-New Zea- 
land, 7; Indian Ocean, about 5; Atlantic Ocean, 7 
(with 5 subspecies of C. coelorhincus); Hawaii, 3; 
eastern Pacific, 6. 

The group, as a whole, is one of upper- 
continental-slope depths. Most species are found 
well above the 1000-meter isopleth, although 
several are known from depths to about 1500 m. 
Only one species, C. labiatus Koehler, 1896, is 
known from below 2000 m (2222 m in the eastern 
Atlantic off Azores; Grey 1956:183). Those 
species that are deepest living and those that live 
in the highest latitudes attain the largest size. 

Depth or habitat preferences are seen among 
some species that are otherwise geographically 
sympatric. Thus in the western Atlantic Ocean, 
Coelorinchus coelorhincus carminatus (Goode, 
1880) is often taken in trawls together with C. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


caribbaeus (Goode and Bean, 1885) and C. occa 
(Goode and Bean, 1885), but depth preferences of 
the three suggest a distinct ‘partitioning of the 
slope bottom with C. caribbaeus shallowest, C. 
coelorhinchus carminatus intermediate, and C. 
occa deepest (Marshall and Iwamoto, in Marshall 
1973). 

In the eastern Pacific the species of Coelorin- 
chus are among the most common fishes in 
upper-slope depths of about 250—1000 m. In some 
areas the fishes are encountered by commercial 
trawlers. C. scaphopsis (Gilbert, 1890) is fre- 
quently captured off Santa Barbara, California, 
by fishermen trawling for Dover sole, Micros- 
tomus pacificus (Lockington, 1879), and 
Pequeno (1971:295) reported finding C. chilensis 
(Gilbert and Thompson, 1916), in the commercial 
catches off San Antonio, Chile. C. fasciatus, 
though not taken in great numbers off eastern 
Pacific shores, is very abundant in other areas. 
Gilchrist (1922:59) has reported trawlers off 
South Africa catching several tons of the fish ina 
single haul. 

The early life history of species of Coelorin- 
chus , as with all other macrourids, is largely un- 
known. Sanzo (1923:125; 1931:56, pl. 5; 1933:255 
et seq. and pl. 16) and Costa (1869:41, pl. 1, fig. 1) 
have reported eggs and larvae of C. coelorhincus. 
Gilchrist’s (1905) capture of the eggs of C. ‘‘fas- 
ciatus’’ in plankton nets attached to beam trawls 
fished on the bottom but not in nets fished simul- 
taneously on the surface suggests that spawning 
in that species takes place near the bottom. Mar- 
shall (1965:318), in providing a tentative life- 
history pattern for macrourids in general, 
suggested that after being shed and fertilized near 
the bottom, the eggs 


develop as they float slowly upward. The larvae hatch at 
levels somewhere near 200 m or continue to rise to these 
levels after emergence. Just before metamorphosis and 
afterwards the young seek deeper waters. Eventually, they 
reach the deep-sea floor. 


Mead et al. (1964:580) added another dimension 
to this pattern in hypothesizing an occasional 
development of expatriate bathypelagic pre- 
juveniles in abyssal species of macrourids. The 
relatively large size (41-44 mm TL) of the four 
juveniles of C. canus (CAS 36797) taken by 
midwater trawl in 0-450 m may indicate that 
their pelagic life was prolonged beyond that 
normal for the species and that they are thus 
expatriates. 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


313 


Figure 4. Ventral view of abdomen of species of 
Coelorinchus showing relative development and placement of 
naked fossa of light organ: (a) Coelorinchus scaphopsis; (b) C. 
fasciatus; (c) C. aconcagua. 


KEY TO THE ADULTS OF EASTERN PACIFIC 
SPECIES OF COELORINCHUS 


la. A prominent, black naked fossa on chest far removed 


Mmombanus\(Higs<4a)) = sok 2 fe Le ee 2 
Ib. A black naked fossa on belly close before anus (Fig. 
4b) fo; mo naked! fossa (Bigs 46) 3 


2a. Head covering thin, mostly transparent. Except for 
ridges, most of head naked or covered with thin, 
nonspinulated scales. Base of tongue black anteriorly 
EE er er ee oe oe ane C. canus (p. 317) 
2b. Head covering rather thick, opaque. Head dorsally 
almost completely covered with coarsely spinulated 
scales. Entire floor of mouth pale 
5 ee ee C. scaphopsis (p. 313) 
3a. Snout short, length much less than diameter of or- 


DE ot ee ae ee 4 
3b. Snout long, length about equal to or (usually) much 
greatemihanidiameten Omonpit aaa eee 5 


4a. A large naked fossa immediately anterior to anus (Fig. 
4b). Gill-rakers 7-11 on medial (inner) side of outer 


iG) Sato ee ee ee eS C. fasciatus (p. 322) 
4b. No fossa on belly (Fig. 4c). Gill-rakers 11-14 on medial 
Sidevomoutemarch==s====a= ana C. aconcagua (p. 319) 


5a. A stout scutelike scale at posterior end of occipital 
region. 7-9 (usually 7 or 8) segmented rays in first 
dorsal fin. Distance between first and second dorsal 
fins less than 2 into orbit diameter. Height of an- 
terior rays of second dorsal less than half height of 
anterior anal rays (Fig. Sa) ______ C. chilensis (p. 332) 

5b. No scutelike scales at posterior end of occipital region. 
9-10 segmented rays in first dorsal fin. Distance 
between first and second dorsal fins more than 2 
into orbit diameter. Height of anterior rays of second 
dorsal more than half height of anterior anal rays 
(Bigs:25b) pee eae eee oe ee C. innotabilis (p. 329) 


SPECIES DESCRIPTIONS 


Coelorinchus scaphopsis (Gilbert) 
(Figures 3F, 4A, 6, 7B, 8) 


Macrurus (Coelorhynchus) scaphopsis Gilbert, 1890:115 (orig- 
inal description; holotype and many paratypes from north- 


FIGURE 5. 


Diagram of (a) Coelorinchus chilensis and (b) 
C. innotabilis comparing presence or absence of stout 
scutelike scale at posterior end of occipital region, relative 
distance between first and second dorsal fins, and relative 
heights of second dorsal fin. 


ern Gulf of California in 265 m; ALBATROsS sta. 3015).— 
Brauer 1906:388 (distribution compiled).—Bohlke 1953:58 
(12 syntypes in Stanford University collection listed). 

Coelorhynchus (Coelorhynchus) scaphopsis: Gilbert and 
Hubbs 1916:144 (listed); 1920:426 (in key). 

Coelorhynchus scaphopsis: Goode and Bean _ 1896:397 
(listed) —Jordan and Evermann 1898:2590-2591 (descrip- 
tion after Gilbert)—Jordan, Evermann, and Clark 1930:207 
(listed) —Lavenberg and Fitch 1966:105 (92 specimens, 
northern Gulf of California) —Makushok 1967:208 (distribu- 
tion compiled).—Chirichigno-F. 1968:416-417 (compared 
with C. canus); 1969:37 (listed). 

Coelorinchus scaphopsis: Iwamoto and Stein 1974:50—S1, figs. 
la, 2a, 3a (distinguishing features; California and Gulf of 
California records). 


DIAGNosis.—A species of Coelorinchus with 
anus slightly removed by 1-3 scale rows from 
anal fin. Long ventral light organ with large an- 
terior dermal window between pelvic bases. 
Subopercle broadly rounded posteroventrally. 
Body moderately deep, greatest depth 5.5-7.5 
into TL. Snout relatively short and blunt, length 
31-35 percent HL, about equal to or (usually) 
slightly less than orbit diameter. Scales with 
spinules arranged in subparallel rows, the middle 
row longest and slightly higher than other rows; 
rows on either side of middle row short, not ex- 
tending to posterior edge of scale; scales present 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


314 


‘Z08LZ SVD Woy UsuTIDads T]-WUI-99 Woy YITIOIO IY S¢sspl SVD JO UY [eSIOP puOdas Jo UISIIO MOJaq BAIR Woy a[BOS :CCCH] SYD JO YUN) pue pesy Jo MaIA [e.QUIA 
‘ZQBLZ PUR 96Z6Z SVD SUdWIDads Wo. spew SUOTIPpR YIM ‘“CoSP] SYO ‘uauoeds TH-WUW-p¢ Woy UMPIP MOIA [RIIIVT “(Waq[ID) sisdoydvos snyouuoja0) “9 AMNOIA 


YAS 
Cy 


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es 


Hf 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


315 


FIGURE 7. 


on lower jaw and underside of head. Mouth 
small, upper jaw 28-32 percent HL. Height first 
dorsal fin about equal to or somewhat longer 
than postrostral length of head. 
DESCRIPTION.—General features as in Figure 
6. Greatest depth of body about 5.5-7.5 into TL. 
Head robust, width over opercles about equal to 
greatest head depth. Trunk and tail moderately 
laterally compressed. Anus slightly removed 
from anal fin by 1-3 scale rows. Head ridges 
stout but not particularly conspicuous; suborbi- 
tal ridge forms an obtuse angle between dorsal 
and ventral head surfaces. Median and lateral 
processes of nasal bones incomplete; anterolat- 
eral margins of snout thus not strongly sup- 
ported by bone. A weak supraoccipital ridge 
formed by a narrow file of 2—4 scales, each with 
high median spinule ridge. Postorbital and 
parietal ridges narrow, fairly sharp. Interopercle 
pointed, without a deep ventral emargination, 
completely hidden beneath preopercle; suboper- 
cle rounded posteroventrally, without a pro- 
duced tip (Fig. 7B). Chin barbel short, thick at 
base but tapering rapidly to filamentous tip. 
Scales of body (Fig. 6) moderate in size, rather 
coarsely covered with subparallel rows of sharp, 
reclined, imbricate spinules; middle spinule row 
longest and highest; 2-4 rows laterally on each 
side of middle row; rows adjacent to middle row 
often incomplete, not extending to edge of scale. 
Head scales (except those forming ridges) cov- 
ered with small spinules in divergent rows. Head 


Right opercle, subopercle and interopercle bones of eight species of Coelorinchus. (A) C. caribbaeus, (B) C. 
scaphopsis; (C) C. canus; (D) C. coelorhincus carminatus; (E) C. aconcagua; (F) C. fasciatus; (G) C. innotabilis; (H) C. 
chilensis . 


surfaces below suborbital ridge covered with 
small loose scales; scaled area extending an- 
teriad as a wedge from preopercle to below an- 
terior margin of orbit. Rami of lower jaws with a 
narrow file of small scales. Gill membranes, ven- 
tral surfaces of snout, lunate areas dorsally be- 
hind anterolateral margins of snout, and area 
around nostrils naked. Terminal snout scute 
broad and blunt. 

Abdominal light organ well developed. A large 
anterior dermal window (length more than half 
diameter of pupil) between and slightly anterior 
to pelvic-fin bases. 

Gill membranes broadly connected to isthmus 
with a very narrow free fold posteriorly. Slits 
between gill arches moderately wide (for genus); 
outer slit about %4 orbit diameter. Uppermost 
(epibranchial) gill-rakers padlike, triangular; 
lowermost raker often triangular, much smaller 
than, and not in alignment with, other rakers; 
middle rakers on each arch more tubercular with 
1-3 sharp, conical, recurved spinules at tip. 

Fins small to moderate in size. Second spi- 
nous ray of first dorsal fin scarcely, if at all, ex- 
tending beyond branched rays; height of first 
dorsal fin usually less than postrostral length of 
head. Second dorsal fin separated by a moderate 
gap from first dorsal fin and weakly devel- 
oped throughout. Pectoral fins extend to, or 
slightly beyond, vertical through anal-fin origin. 
Pelvic fins in larger specimens (more than about 
60 mm HL) fall short of anal-fin origin, but in 


316 


smaller specimens, outer pelvic ray extends be- 
yond that origin. Outer pelvic ray somewhat 
thicker than other rays of fin. 

Intestinal coiling pattern about like that il- 
lustrated by Okamura (1970b: fig. 65B) for 
Coelorinchus asteroides Okamura, 1963; pos- 
terior loop, however, extends anteriad slightly 
farther than shown. Pyloric caeca unbranched, 
slender, relatively long (length about *% orbit di- 
ameter); 15, 20, and 26 in three specimens. Ova- 
ries large in specimens (CAS 29296) taken in late 
October from off Santa Barbara County, Califor- 
nia; individual eggs well formed but small, largest 
ones slightly more than 1.0 mm in diameter. 

Coloration in alcohol generally swarthy with 
bluish to purplish tinge over abdomen, gill cov- 
ers, and gill membranes. A large black lunate 
area behind pectoral-fin bases. Median and pec- 
toral fins generally dusky; pelvic fins blackish, 
but outer ray of fin generally paler. Oral cavity 
completely pallid. Gill cavity generally black ex- 
cept for pale inner region of opercular wall. Gill 
arches somewhat dusky, but rakers and fila- 
ments pallid. Peritoneal lining moderately to 
heavily peppered with large melanophores; 
stomach blackish. 

MEASUREMENTS.—Total length 163+ to 340 
mm, head length 40-80 mm. The following in 
percent of head length [range (¢; n; SD)]: snout 
length 30.5-35.4 (32.33; 19; 1.373); preoral 
length 24.9-32.7 (29.87; 19; 1.776); internasal 
width 19.0—22.8 (21.48; 19; 1.182); orbit diameter 
31.1-36.5 (33.91; 19; 1.545); interorbital width 
21.5-—25.2 (22.98; 19; 1.108); postorbital length 
29:6—33:5 (1.34; 17; .1.331); orbit to angle 
preopercle 33.1—37.2 (35.29; 19; 1.249); suborbi- 
tal width 12.2—14.5 (13.21; 19; 0.568); upper jaw 
length 27.8—32.1 (29.66; 19; 1.224); barbel length 
6.5-10.6 (8.61; 19; 0.852); outer gill-slit length 
16.7-21.1 (19.53; 18; 1.079); preanal length 
132.4-163.8 (143.6; 19; 10.490); outer pelvic to 
anal 33.9—58.9 (43.58; 18; 6.705); isthmus to anal 
57.9-84.9 (68.12; 17; 7.294); greatest body depth 
54.8-72.1 (62.58; 19; 5.147); 1D.—2D. interspace 
21.4-41.7 (33.44; 19; 4.969); 1D. height 55.8-71.7 
(62.13; 18; 4.544); pectoral fin length 43.0-52.5 
(46.59; 19; 2.454); pelvic fin length 32.1-49.6 
(38.67; 18; 4.972). 

Counts.—1D. II, 7-9 (usually 8); pect. i17- 
i119 (rarely 116); inner (medial) gill-rakers, first 
arch 1-2 + 8-10; second arch 1-2 + 7-10; 
scales below 1D. 5-7; below 2D. 4-62; below 
mid-1D. 32-44; over distance equal to predor- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


40 120 ike) 100 30 80 ee 40 
DAS 
ea - 
30} 4+ 
20 
lO; 
O 
10 
20 
30h 
- scaphopsis 
. aconcagua 
40;}— . Canus 
. chilensis 
- fasciatus 
50 
20. l0 100 EON © 1 
FIGURE 8. Map showing eastern Pacific distributions of 


five species of the genus Coelorinchus. 


sal length 27-33 (¢ = 29.81; n = 16); pyloric 
Gaecarls=261(n9— 33): 

COMPARISONS.—Of the eastern Pacific spe- 
cies, C. scaphopsis is most closely related to 
C. canus, but the two are readily distinguished 
by the absence in scaphopsis of a deep notch on 
the ventral margin of the interopercle (Fig. 7b) 
and their different scale spinulation. The former 
feature and the general physiognomy of the head 
and body suggest a closer relationship of 
scaphopsis to C. coelorhincus than to any other 
member of the genus. Scale spinules in C. 
coelorhincus are, however, in a more-or-less 
quincunx pattern in contrast to the distinct rows 
in C. scaphopsis. Although Gilbert and Hubbs 
(1920:426) placed C. scaphopsis close to C. 
patagoniae Gilbert and Thompson, 1916 (=C. 
fasciatus), the significant differences between 
the two in head squamation and _ light-organ 
structure suggest a more distant relationship. 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


7 


SUF 


FiGuRE 9. Coelorinchus canus (Garman). Redrawn and slightly modified illustration from Garman (1899:pl. 84, fig. 1); right 
otolith from a specimen 38 mm in head length (scale line below otolith represents 5 mm); illustration of a scale taken from below 


origin of second dorsal fin of a specimen 35 mm in head length. 


DISTRIBUTION.—C. scaphopsis is known only 
from the northern Gulf of California and off the 
coast of southern California (Fig. 8). Brewer 
(1973:40) reported the species from three cap- 
tures south of the Gulf of California, but exami- 
nation of his specimens (LACM 11748, 13771, 
13778) revealed that they are juveniles of 
Nezumia liolepis (Gilbert, 1890). C. scaphopsis 
is taken occasionally by commercial trawlers out 
of Santa Barbara, California (Rich Lee and Joe 
Copp, personal communication), and Lavenberg 
and Fitch (1966: 105) stated that it ‘‘appears to be 
the commonest rattail in the upper Gulf.’” De- 
spite its apparent abundance in these two areas, 
the authenticated distributional range of C. 
scaphopsis is very limited, perhaps because of 
inadequate sampling. It should be expected in 
outer slope waters of Baja California and along 
most of the mainland coast of Mexico. 

MATERIAL EXAMINED (more than 100 specimens, 8 lo- 
calities). CALIFORNIA:—SIO 61-194 (1, 80 mm HL, 340 mm 
TL), Santa Barbara Co., off Pt. Conception, 265-274 
m.—CAS 29296 (4, 65-72 HL, +260-340 TL), Santa Barbara 
Co., off Gaviota, 287 m, otter trawl, 23 Oct. 1968—CAS 
29298 (1, 67 HL, 280 TL), Santa Barbara Co., off Naples, 296 
m, otter trawl, 18 Nov. 1968.—SIO 67-267 (1, 59 HL, 230 TL), 
Santa Cruz Is., Pelican Bay, 247 m.—CAS 14555 (1, 54 HL, 
245 TL), Santa Cruz Is., Pelican Bay, 202-274 m.—CAS 27802 
(1, 64 HL, 282 TL), W side Santa Catalina Is., 33°25.3’ 
N, 118°32.5W, 183 m, 7 June 1971. GULF OF CALI- 
FORNIA:—SIO 68-94 (>100 spec.), 29°19.9'N, 113°10.4'W, 
bottom trawl.—USNM 44272 (holotype, 61 HL, 250 TL) 
and CAS-SU 179 (11 paratypes, 41-59 HL, 240+ —260 


TL), 20°19'N, 112°50'W, 265 m, beam trawl, ALBATROSS 
sta. 3015, 24 March 1889.—SIO 73-3 (1, 24 HL, 110 TL), 
28°25'N, 112°22'W, bottom trawl, sta. AH 7206, 12 Nov. 1972. 


Coelorinchus canus (Garman) 

(Figure 3E, 7C, 8, 9) 

Macrurus canus Garman, 1899:217-218, pl. 49, fig. 2, pl. 84, 
fig. 1, 2 (original description; illustration; many specimens 
off Panama, ALBATROSS sta. 3355 in 333 m, sta. 3389 in 384 
m, sta. 3391 in 280 m).—Chirichigno-F. 1968: 415-418, fig. 
12 (description; illustration; 11 specimens, 2 localities off 
northern Peru, 4°48’N, 81°17’W); 1969: 37 (listed from 
Ecuador and possibly Peru); 1974: 315, fig. 621 (in key; 
illustration). 

Macrurus (Coelorhynchus) canus: Brauer 1906: 388 (distribu- 
tion compiled). 

Coelorhynchus (Coelorhynchus) canus: Gilbert and Hubbs 
1916:144 (listed); 1920:426 (in key). 

Coelorhynchus canus: Makushok 1967: 208, Table 18 (com- 
piled distribution; implies range of C. canus extends to Gulf 
of California but no documentation given).—Marshall and 
Iwamoto in Marshall 1973:540 (listed).—Parin and 
Makushok 1973:179 (1 specimen off Ecuador, 228-275 m). 


DIAGNosIs.—A species of Coelorinchus with 
a large dermal window of the light organ situated 
between and slightly anterior to bases of pelvic 
fins. Subopercle broadly rounded posteroven- 
trally; interopercle with a deep emargination. 
Height first dorsal fin usually less than postros- 
tral length of head. Snout of moderate length, 
29-37 percent HL, about equal to orbit diame- 
ter. Body scales with small, slender, conical 
spinules aligned in slightly divergent rows; me- 
dian row not enlarged. Scales on head thin, de- 


318 


ciduous, often without spinules. Loose scales 
present on rami of lower jaws and below subor- 
bital region. 

DESCRIPTION.—Head and body moderately 
compressed, greatest width of head 1.2-1.5 into 
its greatest depth. Head covering thin, transpar- 
ent. Bones of head rather thin, mostly transpar- 
ent or translucent. Head ridges low, not particu- 
larly stout or spiny. Supraoccipital crest high, 
but without a row of scutelike scales. Terminal 
snout scute blunt, wide, more spiny ventrally 
than dorsally. Mouth moderately broad, not no- 
tably restricted laterally. Subopercle broadly 
rounded posteroventrally, without a produced 
tip; interopercle with deep cleft in ventral margin 
(Fig. 7C); posteroventral tip barely exposed be- 
yond ventral margin of preopercle. 

All fins except second dorsal fin moderately 
developed. Outer pelvic ray slightly prolonged, 
extending well past anal-fin origin. Other fins 
without produced rays. Second dorsal fin 
rudimentary over most of length. 

Abdominal light organ well developed. Large 
dermal window between and slightly anterior to 
bases of pelvic fins. 

Alimentary canal simple; intestine S-shaped 
or with a short posterior loop before entering 
rectum. Pyloric caeca short, unbranched, 10-14. 
Anus immediately in advance of anal fin, but 
sometimes with one or two scales separating the 
two. 

Head scales, except on ridges, thin and often 
without spinules. Specimens smaller than about 
40 mm HL have broader naked areas and thin- 
ner, less spinulated scales on head. Dorsal and 
ventral surfaces of snout mostly naked. Area 
below ventral surfaces of head scaled from 
below midorbit posteriad to hind margin of 
preopercle. A row of narrow, spinuleless scales 
generally present on rami of lower jaw in 
larger specimens. Body uniformly covered with 
moderate-size spinulated scales. On larger 
scales below dorsal fins of 50-mm-HL specimen, 
scales have 6~7 slightly divergent rows of slen- 
der, conical spinules with up to 8 spinules in 
median row, fewer in lateral rows. Scales on 
pectoral-fin base and shoulder girdle small and 
spinuleless. 

Gill-rakers tubercular, usually two or three 
epibranchial rakers, 7—9 ceratobranchial and 
hypobranchial rakers. Often one or more rakers 
at either end of gill arch very small and platelike. 
Other rakers with one or few short spinules. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


Gas bladder with 6—11 short, flattened retia, 
each connected to a round to kidney-shaped gas 
gland (Fig. 3E). Generally the fewer the number, 
the larger each rete-gas gland complex. All 
specimens in which sex could be macroscop- 
ically determined were females; no drumming 
muscles were present in these. 

Dentition in both upper and lower jaws con- 
sists of small, conical teeth in narrow tapering 
bands. No series of enlarged teeth. 

Coloration in alcohol. Trunk and tail some- 
what flesh colored to swarthy, darker dorsally, 
violet over abdomen, blackish over chest. Gill 
cover and ventral surface of head somewhat sil- 
very in very fresh specimens, but silver color 
lost in long-preserved specimens. Leading edge 
of snout, medial side of pectoral-fin base, margin 
of the first dorsal-fin base black. Rami of lower 
jaw heavily punctate; ventral surface of snout, 
ventral edge of suborbital region, and preopercle 
variously punctate. Large melanophores widely 
scattered over most other parts of head cover- 
ing. Outer edges of gill cavity black; inner sur- 
face pale with scattered melanophores. Outer 
edge of preopercle with heavy punctation. Oral 
cavity mostly pale but blackish along base of 
tongue. Abdominal cavity pale but peppered 
with large melanophores. 

MEASUREMENTS.—Total length 106+ to 234 
mm; head length 30. 1—55.3 mm. The following in 
percent of head length [range (t; 1; SD)]: snout 
length 29.0-37.3 (33.78; 52; 1.602); preoral 
length 27.7-36.0 (31.19; 48; 1.557); internasal 
width 19.4—27.2 (22.12; 50; 1.520); orbit diameter 
29.7-34.8 (32.05; 49; 1.218); interorbital width 
22.9-33.0 (27.64; 48; 1.881); postorbital length 
29.5=35.8 (63.32: 50;) 1.377); orbitetomanele 
preopercle 35.1—40.5 (36.79; 51; 5.324); suborbi- 
tal width 9.3-13.3 (10.89; 49; 0.831); upper jaw 
length 24.4—30.9 (27.28; 50; 1.408); barbel length 
3.2-8.8 (5.99; 49; 1.180); outer gill-slit length 
17.0-21.0 (18.94; 51; 1.009); preanal length 
109.2—133.8 (124.07; 48; 5.208); outer pelvic to 
anal 21.0-38.0 (30.36; 49; 3.336); isthmus to anal 
38.8—61.2 (52.55; 47; 4.051); greatest body depth 
53.0-71.6 (61.47; 48; 4.145); 1D.—2D. interspace 
19.3-38.2 (27.45; 51; 4.649); 1D. height 51.1- 
71.3 (61.52; 43; 5.326); pectoral fin length 42.6— 
60.2 (51.65; 48; 4.594); pelvic fin length 36.4— 
48.8 (42.10; 47; 3.731). 

Counts.—I1D. H, 8-10; pect. 18-24; inner 
(medial) gill-rakers, first arch 1-4 + 8-11 (usu- 
ally 2 + 9-10); second arch 1-3 + 8-11 (usually 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


2-3 + 9-10); scales over distance equal to pre- 
dorsal length 32-36; pyloric caeca 10-14 
(a = IS). 

COMPARISONS.—C. canus appears to be most 
closely related to C. caribbaeus from the west- 
ern Atlantic, sharing with that species a similar 
head physiognomy, large dermal window on 
chest, thin head covering, deeply emarginate in- 
teropercle, rounded posteroventral edge of sub- 
opercle, and rather weakly armed head ridges. 
The two species are readily distinguishable by 
differences in scale spinulation (spinules more 
numerous and not in well-defined rows in carib- 
baeus), orbit diameter (27-31, usually 27-29 
percent HL in caribbaeus , 30-35 in canus), bar- 
bel length (8-11 percent HL in caribbaeus , 3-9 
in canus), and lining of oral cavity (completely 
pallid in caribbaeus, blackish at base of tongue 
in canus). Of the eastern Pacific species, C. 
canus is closest to C. scaphopsis; the two are 
compared in the description of the latter. 

DISTRIBUTION.—The species is known only 
from tropical waters of the eastern Pacific be- 
tween northern Peru (6°20'S) and Costa Rica (to 
about latitude 9°N) (Fig. 8) in depths ranging 
70-360 m. The species might be expected to 
range farther northward to the offings of the 
mouth of the Gulf of California. 

BIOLOGICAL NoTEs.—C. canus is apparently 
bathypelagic during its early life, judging from 
four juveniles (CAS 36797) taken by midwater 
trawl in 0-450 m off the coast of Ecuador. This 
would agree with Marshall’s (1965) ideas regard- 
ing the early life history of slope-dwelling mac- 
rourids. 

Examination of the stomach contents of sev- 
eral mature individuals revealed a preponder- 
ance of copepods and other small crustaceans in 
their diets and the absence of strictly bottom- 
dwelling invertebrates. 

MATERIAL EXAMINED (153 specimens, 22 localities). 
COSTA RICA: CAS 35910-35920, collected by F. H. Berry, 
Dec. 1973 and Mar. 1974, in 192-296 m, 36 specimens (33-43 
mm HL, 111-164 mm TL). PANAMA: GCRL 14250-14256, 
CANopus collections, 1973-1974, in 108-293 m, 15 specimens 
(30-49 HL, 85+ —198 TL). ECUADOR: CAS 35921 (1, 54 
HL, 213 TL), Gulf of Guayaquil, 02°14’S, 81°11.5'W, 120-140 
m, TE VEGA cr. 19, sta. 145, 31 Aug. 1968.—CAS 36797 (4 
juv., 41-44 TL), 03°41'-36'S, 81°36'-20'W, 0-450 m, midwa- 
ter trawl, ANTON BRUUN cr. 16, sta. 623A, 30 May 1966.— 
CAS 35925 (11, 37+ —48 HL, 166-199 TL), 03°50’S, 81°08'W, 
300-360 m, 72-ft otter trawl, ANTON BRUUN cr. 18B, sta. 767 
(LWK 66-116). PERU: CAS 35924 (6, 31-43 HL, 120+— 


168+ TL), 04°57’'S, 81°24'W, 118-133 m, otter trawl, ANTON 
BRUUN cr. 16, sta. 625A, 2 June 1966.—CAS 36802 (1, 42.4 


319 


HL, 192 TL), 04°59’S, 81°27'W, 365-457 m, otter trawl, 
ANTON BRUUN cr. 16, sta. 626B, 3 June 1966.—CAS 35923 
(67, 25-55 HL, 105-234 TL), 05°02'S, 81°24’W, 192-311 m, 
otter trawl, ANTON BRUUN cr. 16, sta. 627A, 3 June 1966.— 
CAS 35922 (10, 21.4-38 HL, 102-159 TL), 06°20’S, 81°01’ W, 
146 m, otter trawl, ANTON BRUUN cr. 16, sta. 633A, 4 June 
1966. 


Coelorinchus aconcagua new species 

(Figures 3D, 7E, 8, 10) 

Coelorhynchus patagoniae: Pequeno, 1971:283—386, figs. 8-10 
(not of Gilbert and Thompson, 1916) (description and figures 
after Gilbert and Thompson, in Thompson 1916; 89 speci- 


mens, Pacific coast Chile, between 30°06'01"S and 
41°43'05"S, 175-428 m). 


DiaGnosis.—A Coelorinchus with a small 
ventral light organ (length less than half pupil 
diameter in adults) situated immediately before 
anus and without an external naked fossa. Sub- 
opercle with a blunt, ventrally pointed tip. 
Height first dorsal fin usually slightly less than 
postrostral length of head. Snout short, 24.2- 
30.3 percent HL, length much less than orbit 
diameter, which is 37.5—43.5 percent HL. Mouth 
relatively large, upper jaw 29-36 percent HL. 
Ventral surfaces of head naked; head ridges all 
relatively low and narrow. Most body scales 
with numerous low, parallel rows of short 
spinules; exposed field of scales often with 
broad spinuleless margins. Gill-rakers numer- 
ous, 11-14 in inner series of first (outer) arch. 

DESCRIPTION OF HOLOTYPE (comments on 
paratypes in parentheses).—General shape as in 
Figure 10. Head large, about five into TL, 
slightly compressed laterally, greatest width 
about 1.3 (1.2—-1.5) into greatest depth. Head 
covering relatively thin, translucent in most 
areas. Supraoccipital crest prominent, forming a 
low hump in dorsal profile. Trunk moderately 
compressed, greatest width over pectoral bases 
about 1.5 (1.4—1.8) into greatest body depth. Or- 
bits huge, forming deep concavities in roof of 
skull. Mouth relatively wide and lateral for a 
Coelorinchus and little restricted laterally; upper 
jaw extends posteriad to below hind third of or- 
bits. Vertical and horizontal margins of preoper- 
cle form broadly acute angle posteroventrally, 
partially covering subopercle, completely hiding 
interopercle; the last two bones shaped as in 
Figure 7E. Ridges of head low and narrow; ter- 
minal and lateral snout scutes weak. Suborbital 
ridge relatively broad behind orbits but narrow 
anteriorly and slightly discontinuous below nos- 
trils; not supported by bone across anterolateral 


320 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


3 ee es 
E a ees os SUN 
: ail ‘i 


Ficure 10. Coelorinchus aconcagua Iwamoto, new species. Lateral view, ventral view of head and trunk, and scale from 
above lateral line below interspace between first and second dorsal fins of holotype, CAS 36779, taken off Chile in 240-260 m. 
Otolith from right side of a paratype, CAS 36800 (56 mm HL). Scale line below otolith represents 5 mm; scale line below tail of 


holotype represents 25 mm. 


margins. Opercular openings rather wide; bran- 
chiostegal membranes moderately restricted, 
with a narrow posterior free fold across isthmus. 
Gill filaments long; gill-rakers tubercular with 
small, fine teeth. 

Premaxillary and dentary dentition consist of 
very small, fine teeth in moderately wide bands 
which extend, in both jaws, to posterior angles 
of mouth. 

Scales relatively deciduous, those on body 
lost. (In larger paratypes, exposed fields of body 
scales with numerous parallel rows of small, 
short spinules, 7-12 rows on largest scales below 
dorsal-fin interspace; broad margins without 
spinules on exposed fields in most body scales. 
Median row of spinules not enlarged except on 
some head scales. Scales on head and chest gen- 
erally with somewhat divergent spinule rows; 
those on chest somewhat coarser than those on 
trunk and tail, and more densely packed.) Ventral 
surfaces of head below suborbital ridge entirely 


naked. Large areas around nostrils and on dorsal 
surface of snout naked. In the holotype (as in 
most specimens), no scales flank the median 
rostral ridge (in some paratypes, a row of thin, 
unarmed scales flank each side of that ridge). 

First dorsal and pectoral fins large, but neither 
longer than postrostral length of head. First 
spinous ray of dorsal fin thornlike; second spi- 
nous ray long but scarcely, if at all, produced be- 
yond adjacent branched rays. (A few small, low 
spinules near tip on leading edge of second spi- 
nous ray in some paratypes.) Second dorsal fin 
low, following rather close behind first dorsal 
fin, poorly developed over most of length, much 
less developed than anal fin. 

Coloration in alcohol brownish or swarthy 
over head; trunk bluish to violet; tail brownish 
with violet tinge. All fins dusky to blackish. Up- 
permost edge of pectoral fin and membrane be- 
tween second spinous and first branched dorsal 
rays black. A prominent blackish area behind 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


pectoral bases. Gill membranes, lips, and chin 
barbel blackish. Oral cavity completely pallid; 
branchial cavity black on medial wall and outer 
margin of lateral wall but pallid on inner portion 
of lateral wall; peritoneal lining blackish but 
peppered with small melanophores. Gill arches 
and rakers blackish, but filaments pallid. The fol- 
lowing information from paratypes: Gas bladder 
well developed, with four large gas glands, each 
connected to a short, broad rete (Fig. 3D). In 
a 52-mm-HL female, anterior bladder wall has 
a thin tough membrane with a slender bundle 
of muscles on each side. A 59-mm-HL male has 
broad sheets of muscles covering anterior end of 
bladder. 

Intestinal coiling relatively simple, about like 
that illustrated by Okamura (1970b: fig. 64B) for 
C. hubbsi Matsubara, 1936. Pyloric caeca slen- 
der, unbranched, lengths about equal to pupil 
diameter; 16, 18, and 20 in three specimens. Ten 
females examined had large ovaries containing 
distinct eggs in various stages of development; 
largest eggs more than 1.0 mm in diameter. 

Light organ in 59-mm-HL paratype small, 
black, flattened, spatulate, lying on epidermis 
within abdominal body wall between abdominal 
muscles, projecting anteriad from base of rec- 
tum, almost reaching pelvic girdle; externally 
apparent only as a blackish area anterior to anus; 
length (7.5 mm) about % of pupil diameter. 

MEASUREMENTS.—Total length 78+ to 345 
mm, head length 19-74 mm. The following in 
percent of head length [range (¢; n; SD)]: snout 
length 24.2-30.3 (26.47; 34; 1.584); preoral 
length 22.8-28.1 (24.19; 34; 1.437); internasal 
width 20.4—25.0 (22.70; 30; 1.162); orbit diameter 
37.4-43.5 (40.39; 34; 1.502); interorbital width 
17.7-23.2 (19.62; 32; 1.356); postorbital length 
2910=34-7,. (Bile73:. 34;) 1.543); orbit to: angle 
preopercle 32.9-36.7 (34.41; 34; 1.015); suborbi- 
tal width 9.0-12.4 (10.20; 33; 0.691); upper jaw 
length 29.2—35.6 (31.68; 33; 1.316); barbel length 
6.2-10.8 (8.56; 32; 1.102); outer gill-slit length 
16.6—23.1 (19.93; 33; 1.431); preanal length 
115.5—139.0 (127.39; 34; 5.722); outer pelvic to 
anal 22.5—40.7 (29.86; 34; 4.128); isthmus to anal 
50.4-74.6 (61.09; 34; 6.113); greatest body depth 
51.8-70.6 (61.83; 30; 5.086); 1D.—2D. interspace 
20.0-46.3 (33.12; 34; 6.240); 1D. height 55.4- 
72.9 (64.03; 24; 4.850); pectoral fin length 51.9- 
66.3 (60.35; 33; 3.246); pelvic fin length 35.8- 
70.5 (45.34; 34; 8.288). 

Counts.—I1D. II, 9-10 (rarely 11); pect. 116- 


321 
i20 (usually i17-i19); gill-rakers, first arch 
2-3 + 9-12; second arch 1-3 + 8-10; scales 


below 1D. 54-7; below 2D. 4%2-7; below mid— 
ID. 3%-4%; over distance equal to predorsal 
length 31-36; pyloric caeca 16-20 (3 specimens). 

COMPARISONS.—In the eastern Pacific the 
new species is likely to be confused only with C. 
fasciatus with which it shares a short snout, 
huge orbit, naked ventral head surface, and rela- 
tively low head ridges. The two are easily differ- 
entiated by the presence in fasciatus of fewer 
gill-rakers (7-9 on first arch vs. 11-14), wider 
suborbital space (15-19 percent of head length 
vs. 9-12), and large naked fossa anterior to anus 
(compared with none in aconcagua). 

Coelorinchus aconcagua shares many fea- 
tures with C. oliverianus Phillipps, 1927, from 
New Zealand waters. The two species have a 
similar physiognomy, huge orbits, short snout, 
thin head covering, relatively large mouth, wide 
gill openings, relatively numerous gill-rakers, 
and relatively weak and narrow suborbital and 
other head ridges. The species are readily dis- 
tinguished, however, by the presence in C. 
oliverianus of a prominent oval naked fossa be- 
tween the pelvic-fin bases (thus suggesting a dis- 
tant relationship), coarser body squamation in 
oliverianus , and a number of morphometric fea- 
tures. 

DISTRIBUTION.—Coelorinchus aconcagua is 
known within a narrow belt bounded by latitudes 
30°06'01"S_ and 41°43'05’S at depths ranging 
175-428 m off the Pacific coast of Chile. Its dis- 
tribution lies somewhat to the north of that of C. 
fasciatus, but the two distributions apparently 
overlap at the 41st parallel. Pequeno (1971:275- 
276) listed four captures (presumably) of this 
species and three of C. fasciatus at that latitude; 
the two species were taken together at two of 
these localities. 

REMARKS.—I have examined five speci- 
mens reported by Pequeno (1971:283) as C. 
patagoniae—four of these were conspecific with 
the new species (one small individual, no. 
P.5.423, was a C. fasciatus), and there is little 
doubt that most others he reported under that 
name are aconcagua. It is a peculiar coincidence 
that Gilbert and Thompson (1916) should de- 
scribe a new Coelorinchus from a single imma- 
ture specimen of C. fasciatus, when an unde- 
scribed species was yet to be found in the same 
area with characters so similar to fasciatus that 
it was later to be confused with that species. 


322 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


FIGURE 11. 


Coelorinchus fasciatus (Ginther). Drawing modified from Giinther (1887:pl. 28, fig. A) of specimen taken from 


east coast of southern extremity of South America, in 256 m, CHALLENGER Sta. 309A. Right otolith from 57-mm-HL specimen 


(CAS 14528). 


ETYMOLOGY.—The specific name is taken 
from the name of the highest peak of the western 
hemisphere, Aconcagua, beneath the shadows 
of which the holotype was captured. The name is 
to be treated as a noun in apposition. 


MATERIAL EXAMINED (97 specimens, 10 localities, all from 
Chile). Holotype:—CAS 36799 (56 mm HL, 269 mm TL), 
33°22'S, 71°53'W, 240-260 m, shrimp trawl, ANTON BRUUN 
cr. 18A, sta. VDM— (field no. LWK 66-18), | Aug. 1966. 
Paratypes:—MNHN, Santiago P.5.463 (1, 69 HL, 296+ TL), 
30°06'01"S, 71°30'06"W, 370 m, CaRLos DARWIN, 27 Apr. 
1963.—LACM 10455-3 (6, 33-48 HL, 150-219 TL), off Val- 
paraiso, 201-119 m, July 1963.—SIO 65-675 (40, 21-68 HL, 
113-305 TL), about 18 miles (29 km) off Valparaiso Harbor, 
22-23 Dec. 1965.—CAS 36801 (19, 40-74 HL, 192-320 TL), 
IMARPE uncat. (2, 46-54 HL, 215-268 TL), 33°22’S, 
71°54'W, 260-280 m, shrimp trawl, ANTON BRUUN cr. 18A, 
sta. 656-0 (field no. LWK 66-16), 31 July 1966.—CAS 36800 
(17, 19-72 HL, 78+ —345 TL), USNM 216705 (5, 40-65 HL, 
200-301 TL) and BMNH 1977.2.4.1-2 (2, 55-61 HL, 230+ — 
301 TL), 35°26’S, 73°01’W, 290-450 m, shrimp trawl, ANTON 
BRUNN cr. 18A, sta. 697 (field no. LWK 66-39), 9 Aug. 
1966.—MNHN, Santiago P.5.393 (1, 74 HL, 320+ TL), 36° 
04'09"S, 73°14'07”W, 340 m, 27 Feb. 1965.—-MNHN, Santiago 
P.5.397 (1, 62 HL, 307+ TL) 39°23’S, 73°50’W, 175 m.— 
MNHN, Santiago P.5.408 (1, 67 HL, 390 TL), 41°11’S, 74° 
12’W, 222 m, 23 Apr. 1966.—MNHN, Santiago P.5.419 (1, 64 
HL, 250+ TL), 41°41'04”S, 74°40’05"W, 250 m, 24 Apr. 1966. 


Coelorinchus fasciatus (Gunther, 1878) 
(Figures 3A, 7F, 8, 11, 12, 13, 14, 15, 16, 17) 


Macrurus fasciatus Ginther, 1878: 24 (original description; 
east coast southern tip South America, CHALLENGER sta. 
305, 309, 311, 73-448 m; erroneously stated as from west 
coast).—Gilchrist 1921:173-174 (eggs from South African 
specimens). 

Macrurus (Coelorhynchus) fasciatus: Ginther 1887:129-130, 
pl. 28, fig. A (redescription of types, CHALLENGER Stations 
corrected to 305A, 309, 309A, 311, 73-448 m; 8 specimens 


illustration).—Brauer 1906:259 (descr.; 16 spec., off South 
Africa). 

Coelorhynchus fasciatus: Goode and Bean 1896:402 (de- 
scription after Giinther)—Garman 1899:397 (listed).—Mc- 
Culloch 1907:348 (1 specimen; 56.3 km E of Sydney, 
Australia; 1463 m).—Gilbert and Thompson, in Thompson 
1916:473 (numerous specimens off SW tip Chile: ALBA- 
TROSS sta. 2783 in 223 m, sta. 278 in 355 m).—McCulloch 
1919:32, pl. 11, fig. 115b (compiled).—Waite 1927:229 
(listed); 1928:5 (listed) —Hart 1946:280 (140 specimens from 
Patagonian Shelf off Argentina).—Poll 1953:228-229, fig. 94 
(32 specimens, off SW Africa, 220 m; description, 
illustration)—Mann 1954:186 (listed from Chile).—Scott 
1970:43 (in key).—Marshall and Iwamoto, in Marshall 
1973:539 (in key). 

Coelorhynchus (Paramacrurus) fasciatus: Gilbert and Hubbs 
1916:144 (name only); 1920:426 (in key).—Barnard 
1925:340-341 (description; South Africa distributions, 163- 
457 m).—McCulloch 1926:177-178 (60 specimens; Bass 
Strait, Tasmania, Great Australian Bight, 268-823 m). 

Coelorhynchus patagoniae Gilbert and Thompson, in 
Thompson 1916:475-476, pl. 6, fig. 2 (original description; 
holotype, USNM 76862, W coast Patagonia between Wel- 
lington Is. and mainland, 355 m).—Mann 1954:186 (listed 
from Chile).—Marshall and Iwamoto, in Marshall 1973:540 
(listed). 

Coelorhynchus (Coelorhynchus) patagoniae: 
Hubbs 1916:144 (listed); 1920:426 (in key). 

Garichthys fasciatus: Whitley 1968:38 (New Zealand records). 


Gilbert and 


DiaGnosis (from South American specimens 
only).—A species of Coelorinchus with a mod- 
erately large light organ (length more than half 
diameter of pupil) situated immediately before 
anus and with a large lenticular dermal window 
extending forward more than half distance to 
pelvic-fin bases. Subopercle produced into an 
acute point. Height first dorsal fin about equal to 
postrostral length of head. Snout short, 29-33 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


23-24 O South America 
@ South Africa 
Fad © South-West Africa 
21-22 & New Zealand (gamma) 
A New Zealand (delta) 
19-20 


NUMBER OF SPINULE ROWS/SCALE 


323 


20-24 24-29 30-34 35-39 40-44 45-49 50-54 55-59 60-64 65-69 70-74 75-79 80-84 85-89 90-94 95-99 


HEAD LENGTH (mm) 


Ficure 12. Scatter diagram showing relationship of head length to number of spinule rows on largest scales of body in five 
populations of Coelorinchus fasciatus. See text for discussion of populations. 


percent HL, much less than huge orbits (39-46 
percent of HL). Mouth small, upper jaw 25-31 
percent HL. Ventral surfaces of head naked; 
head ridges relatively low; body scales with mul- 
tiple, slightly divergent rows of reclined, imbri- 
cate spinules that form distinct low ridges over 
entire exposed field. Scales large, 3 to 4% below 
midbase of first dorsal, usually 21—24 lateral-line 
scales over distance equal to predorsal length. 
Gill-rakers few, 7—9 total in inner series of outer 
arch. 

DESCRIPTION (from South American speci- 
mens; see section on geographic variation for 
discussion of other populations).—Gunther’s 
(1887: pl. 28, fig. A) figure (reproduced in Fig. 
11) is accurate and a good representation of the 
species. Head wide, slightly deeper than wide in 
larger specimens examined, but wider than deep 
in smaller specimens; length about 4.5 into TL. 
Orbits huge, diameter much greater than snout 
length or postorbital length, forming deep con- 
cavities in roof of skull. Snout relatively blunt, 
viewed dorsally or laterally. Mouth relatively 
small, opening somewhat restricted laterally by 


folds of skin; upper jaw extends posteriad about 
to below midorbit. Subopercle with slightly 
pointed posteroventral tip that extends beyond 
posteroventral angle of preopercle. Ridges of 
head relatively low, not prominent except for 
suborbital ridge, which forms a sharp demarca- 
tion between naked ventral portion of head and 
scaled dorsal portion. Anterolateral edges of 
snout not supported by bone. Opercular open- 
ings restricted. Gill membranes broadly attached 
to isthmus, without posterior free fold. Gill fila- 
ments moderately long; rakers very low, tuber- 
cular. 

Premaxillary and dentary dentition consists of 
relatively broad, short bands of cardiform teeth. 
Premaxillary band ending well short of pos- 
terolateral opening of mouth. 

Scales large, strongly adherent on_ thick, 
opaque head covering, somewhat more decidu- 
ous on body. Scale spinules very small (except 
on head ridges), close-packed, reclined, imbri- 
cate, and aligned in sharp, low, ridgelike rows. 
Spinule rows slightly divergent (generally more 
so on head), number of rows varying with size of 


324 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 
PERCENT OF HEAD LENGTH 
SOUTHWEST AFRICA A mm 10 \ 10 B 
SOUTH AFRICA Suborbital Width mn 12 \ _Cmn 12 Snout Length 
SOUTH AMERICA Cm 24 _tn 33 
NEW ZEALAND A M12 \\ =o 12 
NEW ZEALAND yY Gan 9 eee 14 
SOUTHWEST AFRICA | C —m_ 10 
SOUTH AFRICA eaetaeean —_ Cm 12 
SOUTH AMERICA == EE) 9.33 
NEW ZEALAND A —_Cm) 12 
NEW ZEALAND Y cm 14 
SOUTHWEST AFRICA D ML 10 
SOUTH AFRICA Internasal Width = ol 12 
SOUTH AMERICA —OCn_ 33 
NEW ZEALAND A | _tmD 12 
NEW ZEALAND Y a 14 _all 
SOUTHWEST AFRICA E i) 
SOUTH AFRICA Length Upper Jaw m2 
SOUTH AMERICA Ba ole) 
NEW ZEALAND A Cm 12 
NEW ZEALAND Y mn 13 
SOUTHWEST AFRICA F _ (on 10 
SOUTH AFRICA Postorbital Length mim 12 
SOUTH AMERICA a Se 4 
NEW ZEALAND A —Cmlen 12 
NEW ZEALAND y —Omi_ 13 
| 
SOUTHWEST AFRICA G _(min 10 J) 10 H 
SOUTH AFRICA Length Outer Gill Slit Caen 2 Wh ff: Cir) 2 ‘Orbit to Angle 
Z of Preopercle 
SOUTH AMERICA Cm -20 i == EEE] (34 
NEW ZEALAND A nie) Sm 12 
NEW ZEALAND yY ne Tmo 11 /, 14 
JES Se SVs Sa ey) 
FiGure 13. Comparison of eight morphometric features in five populations of Coelorinchus fasciatus using graphic methods 


of Hubbs and Hubbs (1953). See text for discussion of populations. 


scale and size of individual (see Fig. 12). Entire 
ventral surface of head naked; naked areas on 
dorsal surface restricted to nasal fossae and nar- 
row lunate area behind leading edge of snout. 

First dorsal fin long, height about equal to 
postrostral length of head. Spinous second ray 
not produced beyond other rays; leading edge of 
distal portion sometimes armed with a few small 
denticles. Paired fins moderate in length, both 
pectoral and pelvic fins extending beyond verti- 
cal through anus. Outer ray of pelvic fin thick, 
prolonged, extending beyond first several anal 
rays. Anal fin well developed over entire length, 
much higher than second dorsal. 


Coloration in alcohol overall somewhat tawny 
to medium brown. Ventral surfaces of head gen- 
erally paler than remainder of head and body, 
but sometimes peppered with small melano- 
phores. Lining of buccal, branchial, and peri- 
toneal cavities usually blackish. Gill arches 
blackish but filaments pallid. Most fins blackish, 
but pelvic base sometimes whitish. Seven or 
more wide, sometimes rather faint, saddle marks 
on trunk and tail starting from nape. 

Gas bladder (Fig. 3A) very large, bilobed an- 
teriorly with tough, opaque, white, external 
tunic. In a 52-mm-HL female (CAS-SU 23006), 
left lobe contained 4 small retia and gas glands, 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


PERCENT OF HEAD LENGTH 


40 4 50 


5 55 60 65 
SL a ie aL 


linia me 
SOUTHWEST AFRICA A 

SOUTH AFRICA Postrostral length 
SOUTH AMERICA 
NEW ZEALAND /£. 


NEW ZEALAND Y 


70 75 80 
Lal aL) (pl Ly fn FS 


85 90 95 100 
) =} fel SL] [aL LU [LL 


SOUTHWEST AFRICA B 
SOUTH AFRICA Height D, 
SOUTH AMERICA 
NEW ZEALAND A 


NEW ZEALAND yY 


SOUTHWEST AFRICA 
SOUTH AFRICA 
SOUTH AMERICA 
NEW ZEALAND A 


NEW ZEALAND Y 


Length Pectoral Fin 


SOUTHWEST AFRICA 
SOUTH AFRICA 


SOUTH AMERICA cc 


NEW ZEALAND /\ 


NEW ZEALAND Y 


Sees 1 1 


foe SIE Leese 


TTS Js 


(ea eS 


FIGURE 14. 


Comparison of four morphometric features in five populations of Coelorinchus fasciatus using graphic methods 


of Hubbs and Hubbs (1953). See text for discussion of populations. 


right lobe 2 retia and gas glands. In 62-mm-HL 
male from Patagonia (CAS 14528), left and right 
lobes each had two rete-gas gland combinations, 
and posterior chamber had one; large drumming 
muscles present on lateral sides of anterior 
lobes. In two other specimens from New Zea- 
land waters (LACM 11238, 65 mm HL, and 
LACM 10968-13, 60 mm HL) each anterior lobe 
had 2 rete-gas gland combinations and none in 
posterior chamber. 

Coiling of alimentary canal very complex and 
much like that illustrated for C. tokiensis (Stein- 
dachner and Doderlein, 1887) by Okamura 
(1970b: fig. 65C). Pyloric caeca long, slender, un- 
branched, 16, 17, 18, and 19 in four specimens. 

Light organ moderate in size, length longer 
than half pupil diameter; large, lens-shaped der- 
mal window on abdomen immediately anterior 
to anus, falling well short of level of pelvic-fin 
bases. 

MEASUREMENTS.—Summarized and com- 
pared by population in the graphs in Figures 13, 
14, and 15. 

CouNTs (combined for all populations).—1D. 
II, 9-10 (rarely 8 or 11); pect. 115-119 (usually 
116-117); gill-rakers, first arch 7-9; second arch 
7-9 (rarely 10), scales below 1D. 4-512; below 
midbase of 1D. 3-42; below 2D. 3-4%; over 


distance equal to predorsal length of head 20-27 
(usually 23-24). 

GEOGRAPHIC VARIATION.—Examination of 
scattered collections from the South Atlantic 
and South Pacific suggests the possible presence 
of four or five distinct populations of this widely 
distributed species. These populations may each 
warrant formal taxonomic recognition based on 
the differences noted below, but such recognition 
is premature until adequate samples of each are 
examined. A detailed study of additional Coelo- 
rinchus material from off New Zealand, south- 
ern Australia, and the southeastern coast of 
Africa is particularly needed. Study of the litera- 
ture reporting Coelorinchus species from the 
western South Pacific suggests the likelihood of 
closely related Australian-New Zealand species 
having been confused in the past with C. fasci- 
atus. Scott (1970:42-43), in his key to the Aus- 
tralian and New Zealand species of Coelorin- 
chus, appears to have correctly recognized the 
valid nominal species of that region and the 
principal features that characterize each, but it 
is obvious that he worked primarily from the 
literature and not from extensive study material. 
The subtle but consistent differences I have 
found in populations of C. fasciatus from off 
New Zealand are not likely to be recognized in 


326 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 
PERCENT OF HEAD LENGTH 
0 Lies) Gal ea = Tal a uses | ima le a TT PP XT ee Soe — aaa Taal: 250 
SOUTHWEST AFRICA A mn 10 ny 
SOUTH AFRICA Interorbital Width Cm 2 Xx Orbit 
SOUTH AMERICA os iN “eas 
NEW ZEALAND 4 _cme 12 
NEW ZEALAND Y cme 14 
SOUTHWEST AFRICA (e clan 10 
SOUTH AFRICA min 12 Length Posterior Nostril 
SOUTH AMERICA _ co 18 
NEW ZEALAND A m5 
NEW ZEALAND Y tun 14 
L 
SOUTHWEST AFRICA D tm 8 
SOUTH AFRICA oer) Length Barbel 
SOUTH AMERICA Cm 26 
NEW ZEALAND A Cm 
NEW ZEALAND Y Cum 4 
Sa SSeS See e Sea eas YS ee ere) Sera ees 
Ficure 15. Comparison of four morphometric features in five populations of Coelorinchus fasciatus using graphic methods 


of Hubbs and Hubbs (1953). See text for discussion of populations. 


studies using limited material from restricted 
areas. 

From study of material currently available to 
me, it appears that the various populations of C. 
fasciatus can be broadly characterized by sev- 
eral morphometric features, as compared in Fig- 
ures 13, 14, and 15 using the graphic methods 
described by Hubbs and Hubbs (1953). It should 
be borne in mind when examining these dia- 
grams that the sample sizes are generally too 
small to give statistically significant compari- 
sons—the graphs are offered, however, to point 
out possible differentiating characters and char- 
acter trends that should be the focus of subse- 
quent investigations. 

Eastern South Atlantic specimens appear to 
vary in a Clinal pattern from South American 
specimens. Figures 13, 14, and 15 show consid- 
erable differences between South-West African 
and South American specimens in: (1) suborbital 
width; (2) snout length; (3) preoral length; (4) 
internasal width; (5) upper jaw length; (6) post- 
orbital length of head; (7) outer gill-slit length; 
(8) length orbit to angle of preopercle; and (9) 
postrostral length of head. In these characters 
there is little if any apparent overlap between the 
two populations. However, the distributions for 
these same characters in South African speci- 
mens fall in an almost perfectly intermediate po- 
sition, overlapping the distributions in the 
characters of both opposing populations. Were 
these South African specimens not so perfectly 


intermediate, one might suspect, even with the 
limited comparative material, that the South- 
West African population is a species distinct 
from the South American C. fasciatus. 
Additional to these differences in mor- 
phometry between South-West African speci- 
mens (Fig. 16) and those from South America 
are differences in the shape of various parts of 
the head and body. Thus the mouth opening is 
much less restricted laterally in South-West Af- 
rican specimens, the opercle-subopercle height 
is deeper (about 1.05—0.85 of orbit cf. about 
0.85—0.65), the posteroventral angle of the sub- 
opercle is rounded (rather than distinctly acute 
and usually flaplike), the profile of the nape is 
generally more strongly arched, and the dorsal 
profile behind the dorsal-fin origin is more angu- 
lar in its descent to the tip of the tail. Saddle 
markings on the body are completely absent in 
the 10 South-West African specimens examined 
and are generally faint in the South American 
specimens, although Gunther's illustration 
(1887; pl. 28, fig. A) of a specimen from ‘“‘the 
east coast of the southern extremity of South 
America’ is boldly marked with saddlelike 
blotches. The anterior dermal window of the 
light organ is mostly covered with scales in the 
South-West African specimens (and _ partially 
covered in South African specimens) but 
broadly exposed in specimens from other areas. 
Again the South African specimens tend to be 
intermediate in these characters, but, in addition, 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


327 


Li UML 


iv Be; 


FiGurE 16. Coelorinchus fasciatus (Giinther), specimen from South-West Africa in 220 m (after Poll 1953:fig. 94). 


they show a measure of distinctness of their own, 
notably in their apparently longer paired fins and 
higher first dorsal fin (Fig. 14) than in any of 
the other populations compared. South African 
specimens I have examined appeared faded and 
lacked any trace of saddle marks on the body, but 
Smith (1953:fig. 236) illustrated a 35.5—cm speci- 
men that had distinct saddlelike blotches on its 
trunk and tail, and Barnard (1925:340-341), in 
calling the species Banded Rat-tail, described the 
coloration as “‘‘usually with a series of dark 
blotches forming irregular cross-bands.”’ 

The two African populations are obviously 
closer related to each other than they are to the 
other populations of C. fasciatus recognized 
here, despite the distinct clinal pattern they show 
in certain characters. African specimens are set 
apart from South American and New Zealand 
specimens in having (1) a longer postrostral 
length of head, (2) a shorter snout, (3) a shorter 
preoral length, (4) a narrower suborbital width, 
(5) a longer upper jaw, (6) a longer outer gill-slit, 
(7) a somewhat longer posterior nostril (than 
South American and New Zealand delta speci- 
mens but not New Zealand gamma specimens), 
(8) a more narrowly exposed anterior dermal 
window of the abdominal light organ, (9) a longer 
rictus, and (10) a somewhat greater opercle- 
subopercle height. 

It seems that the African populations either 
became separated from the more westerly popu- 
lations at an earlier date than the apparent split in 
the New Zealand and South American popula- 
tions, or the degree of isolation was greater. 
Whatever the case, the amount of differentiation 


that has taken place between African and South 
American populations, on the one hand, is much 
greater than that between South American and 
New Zealand populations on the other hand. 

The specimens that I have called New Zealand 
delta scarcely differ from the South American 
specimens of C. fasciatus , although they tend to 
be more boldly marked, with broader and darker 
saddle marks and, in some, a pale stripe over a 
dark, ground color on the posterior portion of the 
anal fin that is missing in their New World coun- 
terpart. Specimens from the two areas are other- 
wise so apparently close that there is little reason 
to consider them anything but populations of the 
same species. 

The New Zealand gamma population is, on the 
other hand, farther removed from the New Zea- 
land delta populations than the later is to the 
South American populations. The gamma speci- 
mens tend to have fainter saddle markings on the 
body, a more lobate preopercle, a smaller barbel 
and larger posterior nostril (Fig. 15, c & d), a 
somewhat shorter pelvic fin and narrower sub- 
orbital space, fewer spinule rows on the scales 
(Fig. 12), and more pyloric caeca (21, 28, 
29, and 30 in 4 specimens cf. 15—19 in other 
populations). Furthermore, most of the speci- 
mens have small patches of scales on the ventral 
surface of the preopercle, the dorsal fin is gener- 
ally black tipped, and the posterior portion of the 
anal has a black stripe over a pale ground. The 
large number of pyloric caeca appears to distin- 
guish this population from all others of the 
species. I have examined too few collections of 
the two forms off New Zealand to say anything 


328 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


FiGureE 17. 
capture sites of specimens examined; closed symbols represent capture sites reported in the literature. 


definite, but it seems that the delta form is found 
along the southern portion of the New Zealand 
plateau while the gamma form is found farther 
north. It will be interesting to examine Australian 
and Tasmanian collections of C. fasciatus to see 
if they fall into either of the New Zealand forms. 
COMPARISONS.—C. fasciatus appears to be 
most closely related to the Australian species C. 
mirus McCulloch, 1926, from which it differs 
primarily in having a larger orbit, shorter barbel, 
different coloration, and fewer spinule ridges on 
the scales (see Scott, 1970, for additional charac- 
ters). It is also fairly close to C. aconcagua from 
Chile, but is readily distinguished from that 
species by characters given in the key and in the 
description of C. aconcagua. The longer snout, 
smaller orbits, shorter first dorsal fin, and fewer 
spinule rows on scales of C. chilensis make that 
species unlikely to be confused with C. fasciatus. 
REMARKS.—Gilbert and Thompson (in 
Thompson 1916:475) described C. patagoniae 
from a single immature specimen of 125 mm TL 
and compared their new species with C. chilen- 
sis, from which it differs in having a shorter 
snout, fewer scales above the lateral line, and 
other features. Surprisingly, they neglected to 
compare their small holotype with the much larg- 
er C. fasciatus specimens they reported from 
the same ALBATROSS station (no. 2784) or those 
from the previous stations (2783). Comparison of 


General distributions of Coelorinchus fasciatus (triangles) and C. innotabilis (circles). Open symbols represent 


these ALBATROSS C. fasciatus specimens with 
the holotype of C. patagoniae leaves no doubt as 
to their conspecificity. The apparent absence of 
saddle marks in the holotype of C. patagoniae is 
not unusual in that most C. fasciatus specimens 
from ALBATROSS stations 2783 and 2784 are badly 
faded, and they also appear to lack saddle marks. 
DISTRIBUTION.—The species has a subantarc- 
tic distribution (Fig. 17), having been taken from 
Australian-New Zealand waters, off both sides of 
the southern extremity of South America, and off 
southern Africa, at depths of 73-823 m. 


MATERIAL EXAMINED. Eastern Pacific off South America 
(31 specimens, 8 localities):—MNHN, Santiago P.5.423 (1, 21 
mm HL, 97 mm TL), 41°40'03”S, 73°40’05’W, 275 m, 24 Apr. 
1966.—MNHN, Santiago 660037 (1, 62 HL), 41°41'04’S, 
74°40'05"W, 250 m, CARLos Darwin, 24 Apr. 1966.—MNHN, 
Santiago uncat. (1, 53 HL), 42°53’00’S, 72°53’00’W, 150 m, 
CARLOS DARWIN, 9 Apr. 1966.—MNHN, Santiago uncat. (1, 73 
HL), 43°00'08"S, 73°00'04”W, 200 m.—USNM 77290 (3, 46-56 
HL, 200+ —242 TL), USNM 76862 (holotype of C. patagoniae 
Gilbert and Thompson, 29 HL, 125 TL), and CAS-SU 23006 (3, 
50-56 HL), 48°41'S, 74°24’W, 355 m, beam trawl, ALBATROSS 
sta. 2784, 8 Feb. 1888 —USNM 77289 (2, 54-62 HL, 215+ — 
280 TL), 51°02'30’S, 74°08'30’W, 223 m, beam trawl, ALBA- 
TROSS sta. 2783, 6 Feb. 1888 —LACM uncat. (10, 42-68 HL, 
180-284 TL), 52°51'S, 74°13'W, 494-552 m, 12-m otter trawl, 
ELTANIN cr. 21, sta. 22A, 7 Jan. 1966.—LACM 11158-4 (8, 
30-60 HL, 153-260 TL), Straits of Magellan, 52°53'—50’S, 
74°05'-10’W, 544 m, 1.5—m Blake trawl, ELTANIN sta. 1605, 1 
Apr. 1966. 

Western South Atlantic (8 specimens, 3 localities): —CAS 
14528 (3, 59-63 HL, 282-309 TL), Argentina, off Patagonia, 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


ies) 
tN 
‘© 


i) iy a oe 


ee oe 


NWA 
Sys 


RIF 
RES wD WB 


ROgaala 
eas 


5 


35) 


We 


FiGureE 18. Coelorinchus innotabilis McCulloch, CAS 38314, 51-mm-HL specimen taken off Chile in 580 m. Scale line 


represents 25 mm. 


Kayo Maru sta. 10.—USNM 103797 (1, 66 HL, 304+ TL), W 
of Falkland Is., 52°23'S, 65°19'W.—LACM 10458-2 (4, 58-76 
HL, 275-330 TL), off Falkland Is., 53°05'-08’S, 59°31'-24'’W, 
512-585 m, 12-m otter trawl, ELTANIN sta. 339, 3 Dec. 1962. 

Western South Pacific (form gamma) (23 specimens, 4 
localities):—LACM 11238 (2, 65-92 HL, 280-440 TL), New 
Zealand, North Is., Bay of Plenty, 37°49’S, 178°50’E, 732 m, 
12-m otter trawl, ELTANIN sta. 1713, 28 May 1966.—LACM 
11490 (1, 62 HL, 290 TL), New Zealand, W of North Is., 
38°18.4’S, 169°23.1'E, 519 m, Blake trawl, ELTANIN sta. 2200, 
31 May 1968.—LACM 11335(1,35 HL, 175 TL), New Zealand, 
W of Cook Strait, 41°32'-31'S, 174°34'-32’'E, 238 m, 1.5-m 
Blake trawl, ELTANIN sta. 1847, 19 Dec. 1966—DLACM 
10968-9 (1, 71 HL, 325 TL) and LACM 10968-13 (10, 58-100 
HL, 240+ —440 TL), E of Chatham Is., 44°00’—03'S, 178°06’— 
09’W, 430 m, 12-m otter trawl, ELTANIN sta. 1398, 29 Nov. 
1964. 

Western South Pacific (form delta) (12 specimens, 2 
localities):—LACM 109771 (3, 58-71 HL, 244+ —293+ TL), 
SW of New Zealand, 51°00’-01’S, 162°01’E, 371 m, 1.5-m 
Blake trawl, ELTANIN Sta. 1411, 8 Feb. 1965 —LACM 11084-1 
(6, 31-41 HL, 144-188 TL) and LACM 11084—2 (3, 42-50 HL, 
185+ —224 TL), New Zealand slope near Campbell Is., 
53°29'—30'S, 169°48’—45.2'E, 595 m, Blake trawl, ELTANIN sta. 
1989, 1 Jan. 1968. 

South-West Africa (10 specimens, 1 locality):—MRAC 
95772-95781 (10, 51-79 HL, 241-360 TL), 83.7 km S by W of 
Fort Rock Point, 19°52’S, 12°20’E, 220 m, Expédition Océano- 
graphique Belge Atlantique Sud, sta. 108, 25 Jan. 1949. 

South Africa (12 specimens, 6 localities): —SAM 12560 (1, 49 
HL, 262 TL), Table Bay, 285 m.—SAM 12562 (5, 58-63 HL, 
255+ —324 TL), Table Bay, 155m.—SAM 12563 (1, 69 HL, 320 
TL), Table Bay, 240 m.—SAM 12570 (3, 19-38 HL, 104-156 


TL), Table Bay, 282 m.—SAM 12576 (1, 48 HL, 200+ TL), 
36°40'S, 21°26’E, 366 m.—SAM 16375 (1, 99 HL, 420+ TL), 
Table Bay, 366 m. 

Coelorinchus innotabilis McCulloch, 1907 

(Figures 5B, 7G, 18, 19) 

Coelorhynchus innotabilis McCulloch, 1907:348-349, pl. 63, 
fig. 2, 2a (original description; illustrations; holotype, 138 
mm ions and one paratype, 110 mm, both from 56 km E of 
Sydney, Australia, 1463 m); 1919: 32 (list), pl. 11, fig. 11Sa 
(illustration); 1926:180—181 (6 specimens, from eastward of 
Bass Strait by *‘Endeavour’’). 


Coelorhynchus (Paramacrurus) innotabilis Gilbert and Hubbs, 
1916: 144 (list); 1920:429 (in key). 


DIAGNosis.—A species of Coelorinchus with 
anus slightly removed from anal fin. Ventral light 
organ small, not generally visible from exterior, 
its length about 0.5—1.0 into pupil diameter. Sub- 
opercle with a narrow ventral point. Body long 
and slender, greatest depth usually 8-12 into TL. 
Snout slender, sharply pointed, length 41—46 per- 
cent HL. Scales with spinules arranged in sharp, 
slightly divergent ridgelike rows but with no en- 
larged median row; no scales on ventral aspects 
of head except along anterolateral margins of 
snout in large specimens. Mouth small, upper jaw 
21-23 percent of HL. First dorsal-fin height less 
than postrostral length of head; interspace be- 


330 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


tween first and second dorsal fins short, less than 
length of base of first dorsal; height of second 
dorsal moderate, anteriorly almost as high as 
anal. 

DESCRIPTION OF EASTERN PACIFIC SPECIMEN 
(CAS 38314, 51 mm HL).—A long slender fish 
with sharply pointed snout. Head about as wide 
as deep. Trunk and tail moderately compressed 
laterally. Anus slightly removed from origin of 
anal fin by a distance about half diameter of pupil. 
Median and lateral processes of nasal bones con- 
nected and forming a complete bridge across an- 
terolateral edges of snout. Head ridges stout, 
sharp; suborbital ridge separates head into dorsal 
and ventral parts. Occipital and postorbital ridges 
formed of sharply serrated, keellike, median 
spinule rows on scales; median rostral, sup- 
ranasal, and supraorbital ridges low, not formed 
of sharp, keellike scales. Suborbital ridge pos- 
terior to area below middle of orbit composed of 
two rows of scutelike scales that form a double- 
crested ridge. Interopercle completely hidden 
behind preopercle; ventral tip of subopercle pro- 
duced into a flexible, pointed tab, its tip slightly 
exposed beyond posteroventral margin of pre- 
opercle. Barbel thin, short, less than diameter 
of pupil. 

Scales on body of moderate size and covered 
with short, reclined spinules, each aligned in 
close longitudinal rows (8 or 9 rows in large trunk 
scales); outer rows on field very slightly diver- 
gent from middle rows. Posteriormost spinules 
extend beyond scale margin. Middle spinule row 
not enlarged. Scales on head generally with more 
divergent spinule rows. Scales dorsally on head 
mostly without spinules. No enlarged scutelike 
scale at posterior end of occipital sensory canal or 
at anterior end of supraoccipital crest. Broad 
areas around nostrils and behind leading an- 
terolateral margin of snout naked. Terminal snout 
scute somewhat arrowhead shaped in dorsal 
view, dorsoventrally flattened, and armed with 
several longitudinal rows of small spinules, the 
rows diverging posteriorly from the anterior 
apex. 

Light organ very small, externally manifested 
only in a blackish median streak before anus. 
Luminescent gland housed within body wall and 
ina flattened, black, bulbous structure connected 
by a short pedicellike tube to rectum. Entire 
length of gland and tube about equal to half pupil 
diameter. 

Gill membranes broadly connected to isthmus 


without a free posterior fold. Gill slits restricted, 
the outermost about as long as pupil diameter. 
Uppermost (epibranchial) gill-rakers padlike; 
lower rakers tubercular and armed with very 
small spines. 

Fins generally small; first dorsal fin shorter 
than postrostral length of head. Long spinous 
second ray of first dorsal fin barely if at all extend- 
ing beyond adjacent segmented rays. Second 
dorsal fin close behind first dorsal fin, their in- 
terspace less than length of base of latter. An- 
teriormost rays of second dorsal fin relatively 
long for a macrourine grenadier, slightly shorter, 
but distinctly slimmer than anterior rays of anal 
fin; fin height diminishes posteriorly. Uppermost 
ray of pectoral fin a slender splint, closely 
adhered to second ray. Outer pelvic ray some- 
what prolonged beyond inner rays and reaching 
to anteriormost |—3 anal rays. Inner rays extend, 
at most, only to anus. 

Intestine multiply looped and somewhat simi- 
lar to that of C. smithi Gilbert and Hubbs, 1920, 
illustrated by Okamura (1970b: fig. 65D). Pyloric 
caeca number not determined for eastern Pacific 
specimen because of everted stomach and con- 
sequent threat of undue damage if dissected. Two 
New Zealand slope specimens had 7 and 9 short, 
unbranched caeca. Eggs in ovaries of eastern 
Pacific specimen well developed, largest about 
0.8 mm in diameter. Gas bladder in a New Zea- 
land specimen had four large retia and gas glands. 

Coloration in alcohol generally light brown, but 
abdomen and operculum blackish, and ventral 
surfaces of head somewhat greyish. Fins all 
blackish or dusky. Oral, branchial, and perito- 
neal linings black. Gill arches blackish, but 
rakers and filaments pallid. 

MEASUREMENTS.—1Total length 136+ —320 
mm, head length 29-82 mm. The following in 
percent of head length, measurements for eastern 
Pacific specimen first, followed by data for west- 
ern Pacific specimens [range (%; 1; SD)]: post- 
rostral length of head 61.3 [S55.7-61.0 (57.46; 13; 
1.69)]; snout length 40.7 [40.1-46.1 (43.49; 13; 
1.76)]; preoral length 42.1 [40.8—45.9 (43.58; 12; 
1.59)]; internasal width 20.7 [19.6—23.1 (20.94; 13; 
1.02)]; orbit diameter 31.6 [27.7—33.7 (31.11; 13; 
2.07)]; interorbital width 20.0 [16.0-21.9 (18.25; 
13; 1.68)]; postorbital length 28.3 [23.7-27.4 
(25.33; 14; 1.02)]; orbit to angle of preopercle 31.2 
[26.1-31.5 (27.81; 14; 1.66)]; suborbital width 
13.8 [11.9-14.5 (13.17; 14; 0.60)]; upper jaw 
length 22.7 [20.3-21.7 (20.87; 11; 0.46)]; barbel 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


length 8.1 [5.0-7.6 (6.69; 11; 1.25)]; outer gill-slit 
length 11.9 [6.7-9.5 (8.11; 14; 0.96)]; preanal 
length 143.3 [137.0-157.3 (145.22; 11; 6.52)]; 
outer pelvic to anal 34.0 [32.6—42.6 (38.61; 14; 
2.84)]; isthmus to anal 61.9 [54.3—70.6 (64.08; 13; 
5.20)]; greatest body depth 49.0 [35.2-S0.5 
(42.11; 13; 4.77)]; depth over anal origin 41.9 
[30.9-43.8 (36.51; 14; 3.94)]; 1D.—2D. interspace 
Hain. 1=18.4 (12:32; 14> 2-37)]; 1D. height 53.4 
[44.0-53.6 (50.14; 8; 3.15)]; pectoral fin length 
45.4 [37.4—45.3 (41.30; 10; 2.88)]; pelvic fin length 
Alea|b5e2—46:6 68:79: 13; 3:35). 

Counts (eastern Pacific specimen given first 
followed by range of counts for western Pacific 
specimens).—1D. II,10 (II,9-10); pect. 21 (17- 
19); gill-rakers, outer arch 2 + 7 (1-2 + 6-7, 
usually 2 + 6); second arch 2 + 7 (1-2 + 6-7); 
scales below 1D. 9 (542-612); below 2D. 7 (5-7); 
below mid-1D. 6% (444-5!4); over distance equal 
to predorsal length 48 (37-40); pyloric caeca (7 
and 9 in two western Pacific specimens). 

COMPARISONS.—The eastern Pacific specimen 
agrees closely with the original description of C. 
innotabilis, which was based on two small 
specimens. It also compares closely with LACM 
specimens from the western South Pacific ex- 
cept for slight differences in pectoral-ray, gill- 
raker, and scale-row counts. The eastern Pacific 
specimen has 9 rows of scales below the first 
dorsal fin origin, 6% below the mid-base of the 
first dorsal, and 48 lateral-line scales over a dis- 
tance equal to the predorsal length. Comparable 
figures for western Pacific specimens were 
5%-6'2, 442-5’ and 37-40. The body appears to 
be slightly deeper, the postorbital, upper jaw and 
the barbel slightly longer, the outer gill slit de- 
cidedly longer, and the interspace between the 
dorsal fins shorter in the eastern Pacific speci- 
men. In most other features, however, it agrees 
with the western Pacific specimens, especially if 
size-related changes are considered. Thus, in 
specimens of C. innotabilis larger than about 65 
mm HL the scales on the head are more strongly 
spinulated than those of smaller specimens; 
scales on the head that are without spinules in 
small specimens are densely covered with 
spinules in large specimens. The double-crested 
posterior part of the suborbital ridge in small 
specimens is a single broad crest in the largest 
specimens, and scales along the anterolateral 
margin of the snout override the edge of the sub- 
orbital ridge to form a lunate scaled area ven- 
trally in the largest specimens, but not the 


331 


smaller ones. In some New Zealand specimens 
the second spinous ray of the first dorsal fin is 
blackish distally. Most western Pacific speci- 
mens have a blackish edge around the entire or- 
bit, while in the eastern Pacific specimen the 
orbit is blackish only anteriorly and dorsally. 
These slight coloration differences are probably 
negligible; I have examined specimens of C. 
coelorhincus carminatus from the western At- 
lantic that showed even greater color variation 
(Marshall and Iwamoto, in Marshall 1973:558), 
and C. fasciatus populations, noted earlier, 
show considerable differences in body markings. 
If additional material from the eastern Pacific 
shows a consistent difference in the characters 
noted above, it may be reasonable to consider 
the population as distinct at the specific or sub- 
specific level from that of the western Pacific, 
but such designation at present is unwarranted. 

In the eastern Pacific, C. innotabilis is likely 
to be confused only with C. chilensis with which 
it shares a long pointed snout, naked ventral 
snout surface, and small ventral light organ. C. 
chilensis , however, has much coaser scale spinu- 
lation, fewer, more divergent rows of spinules 
on the scales (with most scales having an en- 
larged median spinule row), a lower second dor- 
sal fin, a larger space between the dorsal fins, a 
shorter snout (34—40 percent HL compared with 
4046 in innotabilis), and usually fewer seg- 
mented first dorsal fin rays (normally 7-8, occa- 
sionally 9, compared with 9-10 in innotabilis). 

DISTRIBUTION.—C. innotabilis is known from 
both sides of the South Pacific between latitudes 
33°S and 53°S (Fig. 19). Compared with others of 
the genus, it is a species of moderate depths, 
having been taken between 580 and 1463 m. In 
the eastern Pacific it is apparently sympatric 
with C. chilensis , a species with which the single 
known Chilean representative was captured. Al- 
though C. aconcagua is found within the same 
latitudinal coordinates off Chile, that species ap- 
pears to dwell at somewhat shallower depths of 
about 119-450 m. 


MATERIAL EXAMINED (19 specimens, 7 localities). Eastern 
Pacific:—CAS 38314 (1, 51 mm HL, 215 mm TL), off Chile, 
32°17'S, 71°39.5'W, 580 m, 22-m otter trawl, ANTON BRUUN 
cr. I8A, sta. 702 (field no. LWK 66-44), 11 Aug. 1966. West- 
ern Pacific:—LACM 11238 (4, 34-43 HL, 182-297 TL), New 
Zealand, North Is., Bay of Plenty, 37°49’S, 178°50’E, 732 m, 
12-m otter trawl, ELTANIN sta. 1713, 28 May 1966.—LACM 
11241-5 (5, 29-51 HL, 136-198 TL) and LACM 11241-7 (3, 
52-54 HL, 198-203+ TL), off New Zealand, 38°27’-30'S, 
168°07'-04'W, 659 m, 3-m Blake trawl, ELTANIN sta. 1718, 


332 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


FiGure 19. Coelorinchus chilensis Gilbert and Thompson. Illustration of scale and lateral view of fish from Thompson 
(1916:pl. 6, fig. 1). Scale line below anal fin represents 25.4 mm. Drawings of right otoliths from CAS 38319 (43 mm HL) (top), 
CAS 39319 (71 mm HL) (middle), and CAS uncat. (95 mm HL) (bottom). Scale line below otoliths represents 5 mm. 


12-13 July 1966—LACM 11312-2 (2, 60-62 HL, 237- 
260+ TL), New Zealand, E of Cook Strait, 40°15'-17’S, 
168°16’-18'E, 915 m, 3-m Blake trawl, ELTANIN sta. 1818, 2 
Dec. 1966.—LACM 11485-1 (1, 67 HL, 280+ TL), New Zea- 
land slope, E of South Is., 43°48.2’S, 174°24’-26.3’E, 909 m, 
ELTANIN Sta. 2198, 12 May 1968.—LACM 11449-2 (1, 74 HL, 
320 TL), S of Tasmania, 47°11'10’S, 147°47'-46’E, 1034 m, 
3-m Blake trawl, ELTANIN sta. 1983, 24 Feb. 1967.—LACM 
11085-1 (2, 51-82 HL, 250-300 TL), SW of Campbell Is., 
§3°49’—52.2'S, 169°57.2'—56.1'E, 971 m, Blake trawl, ELTANIN 
sta. 1990, 1 Jan. 1968. 


Coelorinchus chilensis Gilbert and Thompson, 

1916 

(Figures 3C, 5A, 7H, 8, 19, 20B) 

Coelorhynchus chilensis Gilbert and Thompson, in Thompson 
1916:473-474, pl. 6, fig. 1 (original description; illustration; 
holotype and numerous paratypes from ALBATROSS sta. 
2791 off Lota, Chile, in 1238 m).—Makushok 1967:209, table 
18 (depth: distribution)—Pequeno 1971:278—281, fig. 5 (de- 
scription; illustration; 4 specimens, 3 localities, 235-424 m). 


Coelorhynchus (Oxymacrurus) chilensis: Gilbert and Hubbs 
1916:145 (listed); 1920:429 (in key). 


DIAGNosIs.—A species of Coelorinchus with 
a small ventral light organ, its length less than 
half pupil diameter. Ventral surfaces of head 
naked. Subopercle bone with a narrow, ventrally 
pointed tip. Dorsal-fin height less than postros- 
tral length of head. Snout sharply pointed, length 
about equal to or longer than orbit diameter. 
Mouth very small, 20-29 percent of HL. Chin 
barbel short, 5-7 percent of HL. Premaxillary 
teeth in short broad bands which do not extend 
to posterior edge of mouth opening. 

General shape as in Figure 19. Head some- 


what depressed; width over preopercles slightly 
more than greatest head depth. Trunk and tail 
laterally compressed; width of tail over anal-fin 
origin slightly less than twice depth over that 
point. Suborbital ridge stout, acute, separating 
the head into two parts, the ventral part rela- 
tively flattened and without scales, the dorsal 
part rounded and heavily scaled. Head ridges 
prominent, heavily reinforced by stout scutelike 
scales. Median rostral ridge on snout low and 
broad, with spinules relatively short and fine. 
Supranarial and supraorbital ridges narrow and 
low; postorbital and supraoccipital ridges nar- 
row, with median spinules on scales in keellike 
row, forming a relatively high, sharp, ridge axis. 
A single, broad, scutelike scale at anteromedial 
end of nape (anterior end of supraoccipital 
crest); another single scale, this one with a me- 
dian keellike ridge, at posterior end of occipital 
sensory canal (over posttemporal bone), just an- 
terior to lateral-line origin (Fig. Sa). Interopercle 
completely covered by preopercle. Subopercle 
naked and ventrally pointed, the flexible tablike 
tip exposed beyond posteroventral margin of 
preopercle. Gill openings restricted; gill mem- 
branes broadly attached to isthmus and without 
free posterior fold. Gill filaments relatively long. 
Gill-rakers tubercular, coarsely armed with 
short, slender, sometimes recurved spines. 
Body scales large and coarsely spinulated, 
with spinules arranged in distinct, slightly diver- 
gent, ridgelike rows. Spinules in each row larger 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


toward posterior margin of exposed field; middle 
row usually highest, with last spinule extending 
beyond margin of scale. Nasal fossa, entire ven- 
tral surface of head, and narrow margins along 
head ridges naked. 

Fins all relatively small. First dorsal fin lack- 
ing any prolonged rays; fin height much less than 
postrostral length of head. Pelvic fins short and 
small, most rays falling well short of anal-fin ori- 
gin, but the slightly prolonged outermost ray 
reaching just beyond that point. 

Gas bladder well developed. The four retia 
mirabilia short, slender, each connecting to a 
small gas gland (Fig. 3C). Males have large 
sheetlike drumming muscles that encase most of 
the anterior third or so of the gas bladder. An- 
teromedial end of bladder slightly concave, with 
resultant bilobed appearance to bladder head. 
Development of anterior lobes slight compared 
to condition in C. fasciatus (Fig. 3A) and C. 
occa (Fig. 3B). 

Intestinal coiling similar to that illustrated for 
C. smithi by Okamura (1970b:fig. 65D) and of a 
type similar to that of most other members of 
genus. Pyloric caeca moderately long, slender, 
unbranched. 

Light organ small and scarcely discernible in 
exterior view; its only visible manifestation 
being a small, slightly swollen, blackish area an- 
terior to anus. 

Overall coloration in alcohol greyish-brown. 
Fins blackish. Oral, branchial, and peritoneal 
membranes blackish. Naked ventral surfaces of 
head whitish to dirty brown. Lips and chin bar- 
bel pallid. 

MEASUREMENTS.—Total length 123-483 mm; 
head length 34.0-102 mm. The following in per- 
cent of head length [range (¢; n; SD)]: snout 
length 34.0-40.1 (37.40; 39; 1.508); preoral 
length 31.0—37.2 (33.89; 39; 1.808); internasal 
width 18.2—22.4 (20.05; 39; 1.042); orbit diameter 
29.0-38.3 (33.94; 39; 2.124); interorbital width 
18.9-25.0 (22.06; 37; 1.529); postorbital length 
25.5-31.1 (28.06; 39; 1.499); orbit to angle of 
preopercle 28.2—36.0 (31.12; 38; 1.718); suborbi- 
tal width 13.3—16.8 (14.65; 37; 0.883); upper jaw 
length 20.0-—29.2 (25.03; 37; 1.977); barbel length 
3.9-10.0 (6.40; 37; 1.184); outer gill-slit length 
9.5-17.7 (12.45; 35; 1.571); preanal length 
131.8-150.4 (140.14; 36; 5.616); outer pelvic to 
anal 23.2-45.0 (30.86; 36; 4.330); isthmus to anal 
47.2-72.5 (55.16; 37; 5.235); greatest body depth 
45.3-60.7 (52.44; 31; 3.892); 1D.—2D. interspace 


333 


FiGureE 20. Dorsal view of nasal bones of (a) Coelorinchus 
occa and (b) C. chilensis comparing relative development of 
median and lateral processes in the two species. 


21.7-36.3 (29.114; 37; 4.564); 1D. height 41.4— 
49.3 (45.30; 39; 2.666); pectoral fin length 38.9- 
50.0 (42.46; 35; 2.621); pelvic fin length 31.3- 
42.0 (38.58; 32; 2.831). 

Counts.—I1D. II, 7-9; pect. 116-121 (usually 
118-120); gill-rakers, first arch 1-2 + 7-9; sec- 
ond arch 1-2 + 6-8, scales below 1D. 4%2-6 
(usually 5%); below 2D. 3'%2-6 (usually 4%); 
below mid-1D. 34-5 (usually 3%); over dis- 
tance equal to predorsal length 33-41 (35.33; 21; 
2.556); pyloric caeca 9-12 (n = 3). 

CoMPARISONS.—Of the eastern Pacific mem- 
bers of the genus, C. chilensis is likely to be 
confused only with C. innotabilis with which it 
shares a long, slender, pointed snout and a 
naked ventral snout surface. It is, however, 
readily distinguished from that species by its 
fewer scale rows below the first and second dor- 
sal fins, longer interspace between the dorsal 
fins, more coarsely spinulated scales, and a 
smaller light organ. The combination of sharply 
pointed snout, naked ventral snout surface, 
coarsely spinulated scales, small light organ, and 
small mouth immediately separates C. chilensis 
from other eastern Pacific congeners. 

REMARKS.—Coelorinchus chilensis was rele- 
gated by Gilbert and Hubbs (1920:429) to the 
subgenus Oxymacrurus, apparently because of 
the relatively low median row of spinules on its 
scales, which contrasts with the very strong 
keellike median row in species of the subgenus 
Oxygadus. But in other characters given in their 
analytical key (anus immediately before anal fin; 
second dorsal spine less than postrostral length 
of head; body without dark markings), C. 
chilensis agrees with both Oxygadus and 
Oxymacrurus. 


ww 
ww 
. 


Okamura (1970a:153—154) used other charac- 
ters besides the size of the median spinule row to 
distinguish the two subgenera. Thus he charac- 
terized Oxymacrurus as having the anterolateral 
margin of the nasal bones entire, the luminous 
organ decidedly longer than half the pupil diame- 
ter, no scute at the posterior end of the occipital 
sensory canal, and median rostral series of 
scales flanked on each side by a single series of 
scales. Oxygadus, in contrast, supposedly has a 
broad gap between the median and lateral pro- 
cesses of the nasal bone (anterolateral margin is 
thus interrupted), a very small luminescent 
organ whose length is less than half the occipital 
sensory canal, and the median rostral series of 
scales flanked on each side by two or three 
series of scales. C. chilensis agrees with Oka- 
mura’s characterization of subgenus Oxygadus 
except for the size of the median spinule row and 
by the presence of only one series of scales 
flanking the median rostral scale row. The 
species thus appears to bridge the gap separating 
the two nominal subgenera, and its placement is 
left in doubt. 

The supposed presence in Oxygadus of a 
broad gap between the median and lateral pro- 
cesses of the nasal bone might seem to be a 
clear-cut distinguishing character, but C. occa,a 
species otherwise closely resembling other 
members of the ‘“‘oxygadus-group,’’ has median 
and lateral processes almost united (Fig. 20). 
Okamura’s use, as a diagnostic character, of the 
number of scale rows flanking the median rostral 
series of scales (and separated by a narrow, 
naked trough from those scales next to the su- 
pranarial ridges) is of questionable value, for he 
himself states that C. (Oxymacrurus) tokiensis 
(Steindachner and Déderlein) has two rows on 
each side (the same as in species of Oxygadus), 
and my examination of two specimens (CAS-SU 
22924) of C. (Oxymacrurus) japonicus (Tem- 
minck and Schlegel) also showed two rows (and, 
in fact, Okamura’s (1970a: text-fig. 78) illustra- 
tion of the head of that species also suggests the 
presence of more than one row on each side). 

Based on these facts, if C. chilensis is in- 
cluded in subgenus Oxygadus, only two unam- 
biguous characters remain for distinguishing 
Oxygadus and Oxymacrurus: 1) the relative size 
of the luminescent organ (length greater than 
half pupil diameter in Oxymacrurus, less than 
half in Oxygadus); and 2) the presence of a 
scutelike scale at the posterior end of the occipi- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


tal sensory canal in Oxygadus that is lacking 
in Oxymacrurus. Including C. chilensis in 
Oxymacrurus leaves only one unambiguous 
character—that of the relative development of 
the median spinule row on the scales. 

Obviously, a more thorough study is needed 
to clarify the relationships within and between 
the subgenera, and to show if they indeed repre- 
sent natural groups; but that is beyond the scope 
of the present work. For now, it seems best to 
forego any subgeneric allocation of C. chilensis. 

DISTRIBUTION.—The species is known from 
the coasts of Peru and Chile between latitudes 
6°S and 38°S, in depths ranging 260-1480 m. The 
1220-m difference between its shallowest and 
deepest captures is considerable for a species of 
Coelorinchus, but that range is not unusual 
for species of other genera (especially Cory- 
Phaenoides and Nezumia). There was no cor- 
relation between latitude and capture depths of 
C. chilensis. 

The species is apparently nowhere found in 
great abundance, although Pequeno (1971:285) 
reported its presence in commercial catches off 
San Antonio, Chile (latitude 33°34’S). The most 
taken in seven ANTON BRUUN stations (at which 
commercial-size shrimp trawls of about 22-m 
head rope were used) was 13 (station 714, in 950 
m). 


MATERIAL EXAMINED (71 specimens, 11 localities). 
Peru:—LACM 33884-1 (1, 330 mm TL), due W of Lobos de 
Tierra, 06°26'S, 80°05’'W, 1025 m, beam trawl, 23 Jan. 1974. 
Chile: —USNM 216706 (5, 51-90 mm HL, 191-335 mm TL) 
and CAS 38319 (8, 50-76 HL, 179-305 TL), 24°29.5’S, 
70°40'W, 950 m, 22-m otter trawl, ANTON BRUUN cr. 18A, 
sta. 714 (LWK 66-60), 16 Aug. 1966.—CAS 38316 (1, 41 HL, 
155 TL), 32°08.5’S, 71°43'W, 960 m, 22-m otter trawl, ANTON 
BRUUN cr. 18A, sta. 703 (LWK 66-47), 12 Aug. 1966.—CAS 
38313 (5, 34-67 HL, 123-240 TL), 32°17’S, 71°39.5'W, 580 m, 
22-m otter trawl, ANTON BRUUN cr. 18A, sta. 702 (LWK 
66-44), 11 Aug. 1966.—CAS 38308 (2, 82-88 HL, 280-280 
TL), 33°22'S, 71°54'W, 260-280 m, shrimp trawl, ANTON 
BRUUN cr. 18A, sta. 623A (LWK 66-16), 31 July 1966.—CAS 
38311 (5S, 61-78 HL, 200-300 TL), 33°39’S, 72°09’W, 1170- 
1480 m, 22-m otter trawl, ANTON BRUUN cr. 18A, sta. 699 
(LWK 66-41), 10 Aug. 1966.—CAS 38309 (13, 48-95 HL, 
195-367 TL), LACM 36019-1 (3, 49-58 HL, 195-210 TL), and 
CAS 38333 (3, 96-102 HL, 445-480 TL), 34°06.5’S, 72°18'W, 
750 m, 22-m otter trawl, ANTON BRUUN cr. 18A, sta. 687 
(LWK 66-25), 5 Aug. 1966.—CAS 38385 (9, 41-66 HL, 160- 
244 TL), 34°53.5'S, 72°44’'W, 780-925 m, 22-m otter trawl, 
ANTON BRUUN cr. I8A, sta. 698 (LWK 66-40), 9-10 Aug. 
1966.—CAS 38349 (3, 90-101 HL, 403-483 TL), 35°26.5’S, 
73°01'W, 290-450 m, 22-m otter trawl, ANTON BRUUN cr. 
18A, sta. 697 (LWK 66-39), 9 Aug. 1966.—USNM 76861 
(holotype, 80 HL, 310 + TL), USNM 76890 (6 paratypes, 
58-76 HL), and CAS-SU 22726 (7 paratypes, in poor condi- 


IWAMOTO: EASTERN PACIFIC MACROURIDS 


tion), off Lota, 38°08’S, 75°53’W, 1238 m, ALBATROSS sta. 
2791, 14 Feb. 1888. 


RELATIONSHIPS OF EASTERN PACIFIC 
COELORINCHUS 


The six eastern Pacific species of Coelorinchus 
each show closer affinities with species outside 
the region than they do with one another. The two 
northernmost species, C. scaphopsis and C. 
canus, are obviously related to three Atlantic 
species, C. coelorhincus (Risso, 1810), C. carib- 
baeus (Goode and Bean, 1885), and C. ventrilux 
Marshall and Iwamoto, 1973. They share in 
common the following features: 1) a well devel- 
oped light organ with large dermal window be- 
tween the pelvic-fin bases; 2) relatively long, 
narrow, tapered bands of teeth in both jaws; 3) 
relatively finely spinulated scales on the head and 
body; 4) head ridges not especially sharp and 
prominent; 5) subopercle terminating ventrally in 
a rounded tab; 6) ventral surfaces of head mostly 
scaled; and 7) gill opening relatively wide. C. 
scaphopsis is closest to C. coelorhincus , whose 
various populations are found throughout the At- 
lantic, while C. canus is closest to C. caribbaeus , 
whose primary distribution lies in the Gulf of 
Mexico and Caribbean Sea. These last two 
species and C. ventrilux have a deep notch in the 
ventral margin of the interopercle (Fig. 7), an 
elongated terminal snout scute, weakly spinu- 
lated head ridges, and thin, transparent head cov- 
ering. C. scaphopsis and C. coelorhincus , on the 
other hand, have no ventral notch in the in- 
teropercle, a blunt, 3-pronged terminal snout 
scute, rather coarsely spinulated head ridges, and 
thick, opaque head covering. 

Of the four other eastern Pacific species, two 
(C. fasciatus and C. innotabilis) are widespread 
southern hemisphere occupants and two (C. 
chilensis and C. aconcagua) are restricted to the 
Pacific coasts of Chile and Peru. C. fasciatus has 
a broad subantarctic distribution, having been 
taken off southern Africa, both sides of the tip of 
South America, off New Zealand, and off south- 
ern Australia. Based on the structure of the ven- 
tral light organ, its affinities appear to lie chiefly 
with other Australian macrourids, most notably 
with C. mirus McCulloch, 1926, but also with C. 
aspercephalus Waite, 1911, C. mortoni Ogilby, 
1897, and C. australis (Richardson, 1839). 

C. innotabilis, which ranges from southern 
Australia to Chile, does not appear to be closely 
related to any other Australian or Chilean 


335 


species. Rather, its affinities are with species of 
the subgenus Paramacrurus (as defined by Gil- 
bert and Hubbs, 1920) from the Philippines, East 
Indies, Japan, and Hawaii, and it seems closest to 
C. cingulatus Gilbert and Hubbs, 1920, from 
northern Luzon and Formosa. C. innotabilis is 
readily distinguished from C. cingulatus, how- 
ever, by the lack of body markings so prominent 
in that species. 

The relationships of C. aconcagua and C. 
chilensis are rather obscure. C. chilensis appears 
to be closest to various Indo-Pacific members of 
the subgenus Oxymacrurus , yet it shares several 
important diagnostic characters of subgenus 
Oxygadus , whose members are found in Hawaii, 
in the western Pacific, the Atlantic, and the In- 
dian Ocean. C. aconcagua shares many features 
with C. oliverianus from New Zealand waters, 
including a similar physiognomy, huge orbits, 
short snout, thin head covering, relatively large 
mouth, wide gill openings, relatively numerous 
gill-rakers, and relatively weak and narrow sub- 
orbital and other head ridges. The large naked 
fossa of the light organ between the pelvic-fin 
bases and the shape of the subopercle in C. 
oliverianus , however, suggest a distant relation- 
ship for the two species. The new species may be 
fairly close to C. fasciatus, but the large fossa 
immediately anterior to the anus in that species 
contrasts with the complete absence of a naked 
ventral fossa in C. aconcagua. 


ACKNOWLEDGMENTS 


I thank the many persons from whom I have 
directly or indirectly received assistance during 
the preparation of this paper. I may have failed 
to remember some in writing this acknowledg- 
ment—for those latter my apologies for the in- 
advertent omission. 

The following curators and their aides assisted 
in the loan or examination of material in their 
care: N. Bahamonde-N. (MNHN, Santiago), C. 
E. Dawson (GCRL), P. A. Hulley (SAM), E. A. 
Lachner and V. G. Springer (NMNH), R. J. 
Lavenberg, M. Ruggles, J. Newman (LACM), 
R. L. Rosenblatt, J. Pulsifer, J. Copp (SIO), C. 
L. Smith and D. E. Rosen (AMNH), D. Thys 
van den Audenaerde (MRAC). R. Lee (formerly 
with Santa Barbara Natural History Museum), 
F. H. Berry, and G. C. Miller provided other 
specimens of Coelorinchus for this study. I give 
special thanks to L. W. Knapp of the Smithson- 
ian Oceanographic Sorting Center for allowing 


336 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 12 


study of the extensive macrourid collections of 
the R/V ANTON Bruun. J. E. Fitch (California 
Fish and Game) provided information on 
otoliths. 

Technical assistance was provided by T. 
Prudhomme (preliminary data gathering), J. E. 
Gordon, W. C. Ruark, K. Hakanson (X-ray and 
curatorial assistance), M. Giles (photography), 
and B. Powell (typing). Katherine P. Smith ren- 
dered the artistic, yet technically precise, illus- 
trations. 

L. Dempster and W. N. Eschmeyer critically 
reviewed the manuscript and offered useful ad- 
vice. 

This research was supported by a grant in aid 
of research from the National Science Founda- 
tion (BMS 7503153) to the author. 


LITERATURE CITED 


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1887. Report on the deep-sea fishes collected by 
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IWAMOTO: EASTERN PACIFIC MACROURIDS 


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337 


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PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 13, pp. 339-343 


February 8, 1978 


THE HARRY E. SOVEREIGN COLLECTION OF 
NORTHWEST PACIFIC DIATOMS 


By 


Albert Mahood 


Associate, Geology Department, California Academy of Sciences, Golden Gate Park, 
San Francisco, California 94118 J 


ABSTRACT: The Harry E. Sovereign collection, now at the California Academy of Sciences, is a significant 
contribution to the knowledge of the recent and fossil diatoms of the Pacific Northwest. The extensive 
collection, the work of over 30 years, represents a wide variety of sampling environments and is especially rich 
in freshwater forms. It contains important historical data concerning the conditions of lakes before the impact of 
current populations, making the collection invaluable to future investigations of the lakes of Washington State 


and adjacent areas. 


INTRODUCTION 


Harry E. Sovereign was born in Denver, Col- 
orado, in 1884. After graduation from the Univer- 
sity of Colorado, in 1908, he began his profes- 
sional career as a civil engineer for the firm of 
Field, Fellow and Hinderlider. From 1911 to 1915 
he worked for the Wilcox Canal Company on the 
construction of a dam near Bear Lake in Idaho. 
For the following fifteen years he was employed 
by the American Metals Company in the devel- 
opment of silver mines in Mexico. In 1927 Mr. 
Sovereign and his wife, Ruby, moved to Seattle, 
Washington, where he worked for the Henry W. 
Beecher Company on naval engineering projects 
until his retirement in 1953. Mr. Sovereign’s in- 
terest in diatoms began shortly after his arrival in 
Seattle. He notes that Professor Trevor Kincaid, 
University of Washington, furnished him with his 
first collections of Pacific Northwest diatoms. 
With this incentive, Sovereign pursued his re- 
search and field collecting on an active basis until 
approximately 1963. In honor of Professor Kin- 
caid, he named a new species of diatoms, 
Navicula Kincaidii, published in the Proceedings 
of the California Academy of Sciences (1963). 

During the last few years of his life Mr. 


Sovereign was ably assisted by Mrs. Sovereign, 
especially in the preparation of an unpublished 
paper, “‘The diatoms of Ohanapecosh Hot 
Springs.’ Harry E. Sovereign died in Seattle on 
April 15, 1965, and his wife died in May of the 
same year. As a bequest of the Sovereign estate, 
the complete collection including his diatom li- 
brary and microscope was given to the California 
Academy of Sciences where it is now a part of the 
diatom collection in the Department of Geology. 

Sovereign’s extensive correspondence with 
Friedrich Hustedt, the well known German au- 
thority on diatoms, is preserved as an integral 
part of the collection. 

Although he published only two papers, the 
taxonomic and environmental value of his collec- 
tion has prompted the writing of this paper. 


DESCRIPTION OF THE COLLECTION 


The collection may be divided into three gen- 
eral areas of interest to the reader. The first and 
most important is composed of recent flora of the 
lakes, streams, and creeks of the State of Wash- 
ington. Sovereign emphasized population varia- 
tions related to environmental differences 
created by the Cascade Range as it divides the 


[339] 


340 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 13 


State of Washington. He collected 452 samples 
from Washington and the neighboring states in 
support of his efforts to confirm the population 
variations. He divided the state by geographic 
and environmental boundaries into five study 
areas: 1) Eastern Washington, including Stevens, 
Pend d Oreille, Spokane, Whitman, and Garfield 
counties; 2) Central Washington, including 
Okanagan, Ferry, Lincoln, Adams, Franklin, 
Walla Walla, Benton, Yakima, Kittitas, Grant, 
Douglas, and Chelan counties; 3) Mount Rainier 
National Park; 4) Western Washington, including 
Whatcom, Skagit, Snohomish, King, Pierce, 
Thurston, Pacific, Lewis, Cowlitz, and Skamania 
counties; 5) The Olympic Peninsula including 
Clallam, Grays Harbor, and Mason counties. 

With the exception of Ferry, Garfield, Benton, 
and Wahkiakam counties, samples were col- 
lected from representative lakes, streams, 
creeks, and occasional snow rills. Sovereign’s 
attention to detail and numerous field notes has 
allowed us to establish township and range or 
longitude and latitude for most samples. 

The second area of interest is fossil com- 
munities from both Washington and Oregon. 
Some of the sample locations have since been 
inundated by subsequent formation of lakes. The 
fossil samples are located by township and range 
as well as field sample numbers. 

The third area of interest encompasses a wide 
range of locations and environments including 
recent, fossil, freshwater, and marine diatoms. 
The samples include specimens from Manila Bay, 
Philippine Islands, Colorado, California, and 
Alaska. 

The collection proper consists of seven units: 
(1) a catalog of the collection, (2) a species list, (3) 
approximately 1400 strew mounts, (4) about 600 
vials of cleaned and preserved material, (5) a card 
index of the collection, (6) field notebooks, (7) 
and an unpublished manuscript, ‘‘The diatoms of 
Ohanapecosh Hot Springs.”’ 

The catalog is a single volume in which samples 
are listed in chronological order and assigned 
consecutive numbers from | to 548. These num- 
bers appear on all subsequent references to a 
particular sample. The number is followed by a 
description of the location, date of sampling, ini- 
tials of the collector, pH, alkalinity, elevation, 
and important environmental data. 

The species list is a series of notebooks con- 
taining lists of species for each sample prepared 
by Sovereign. The list contains the microscope 


coordinates for the location of each species on the 
corresponding numbered slide. In addition, the 
list commonly includes the reference to original 
literature for the species, with occasional draw- 
ings of definitive structures and morphological 
measurements. 

The third unit is a set of cleared, strewn, hyrax 
mounts of these prepared samples. The slides are 
numbered according to the catalog entry. In sev- 
eral cases more than one slide for a particular 
sample was prepared. For example, slides pre- 
pared from sample 234 have 234 on the slide label 
and the first slide is 234-1, the second 234-2, etc. 
These numbers appear on the species list to indi- 
cate which slide was used. 

In the card index samples are cross-indexed by 
species, general location and specific area, such 
as Mount Rainier or Crater Lake. In all three 
categories the samples are indexed by catalog 
numbers. Each species card contains the slide 
numbers on which the species was identified. 

Sovereign’s field notes are found in several of 
the volumes or transposed to volumes of specific 
date; e.g., notes concerning pH are found in a 
single volume. 

To facilitate the use of this collection, I have 
prepared a summary, arranged chronologically 
and cross-indexed by geographical names. It con- 
tains a common number that refers to sample, 
location and slide. The summary also includes 
township and range, or longitude and latitude, 
description of location, county and/or state, date 
of collection, pH and species lists. This summary 
is on file with the Sovereign collection. 


ACKNOWLEDGMENTS 


I gratefully acknowledge the assistance of Mrs. 
Margaret M. Hanna, Research Associate in the 
Geology Department of the California Academy 
of Sciences, for her enthusiastic support with the 
research and development of this manuscript. I 
would also like to thank Peter U. Rodda, Chair- 
man of the Geology Department, for the use of 
the Sovereign collection. 


BIBLIOGRAPHY OF HARRY E. SOVEREIGN 


1934. Diatomite in the State of Washington. Illustrated with 
photographs. Unpublished report made for Washington 
State Geological Survey. 35 pp. 

1952. Diatoms of Ohanapecosh Hot Springs. Unpublished 
manuscript. 8 pp. 

1958. The diatoms of Crater Lake, Oregon. Trans. Am. 
Micro. Soc. 77(2):96—-134. 

1963. New and rare diatoms from Oregon and Washington. 
Proc. Calif. Acad. Sci., Ser. 4, 21(14):349-368, 2 pls. 


MAHOOD: SOVEREIGN DIATOM COLLECTION 


SPECIES AND LOCALITIES LIST FROM SLIDES PREPARED BY FRIEDRICH HUSTEDT 


IN THE H. E. SOVEREIGN COLLECTION 


Species 


Achnanthes brevipes Agardh 
Achnanthes coarctata Brébisson 
Achnanthes grimmei Krasske 
Achnanthes hungarica Grunow 
Achnanthes kryophila Boye Peterson 
Achnanthes lapponica Hustedt 
Achnanthes linearis (Wm. Smith) 
Grunow 
Achnanthes marginulata Grunow 
Achnanthes minutissima Kitzing 
Achnanthes minutissima Kitzing 
Achnanthes pyrenaica Hustedt 
Achnanthes inflata Kitzing 
Achnanthes trinodis Arnott 
Amphora coffeaeformis Agardh 
Amphora coffeaeformis Agardh 
Anomoeoneis styriaca (Grunow) 
Hustedt 
Anomoeoneis zellensis (Grunow) Cleve 
Caloneis clevei Grunow 
Caloneis ladogense (Cleve) Hustedt 
var. densestriata Hustedt 
Caloneis obtusa (Wm. Smith) Cleve 
Cocconeis disculus (Schumann) Cleve 
Cyclotella distinguenda Hustedt 
Cyclotella elgeri Hustedt 
Cyclotella iris Brun 
Cyclotella stelligera Cleve & Grunow 
Cymbella alpina Grunow 
Cymbella aspera (Ehrenberg) Cleve 
Cymbella bernensis Meister 
Cymbella botellus Lagerstedt 


Cymbella cistula var. arctica Lagerstedt 


Cymbella gracilis (Rabenhorst) Cleve 


Cymbella hebridica (Gregory) Grunow 
Cymbella lapponica Grunow 
Cymbella norvegica Grunow 
Cymbella ruttneri Hustedt 


Cymbella sumatrensis Hustedt 
Cymbella turgidula Grunow 


Diatomella balfouriana Greville 


Denticula tenuis Kitzing 


Location 


Kolberg, Salzwiesen 2 

Zittau Granitwand Kr. 1581-2 
Kleinern bei Wildungen 28.8.29.H 
Klinkerteich, Plon 

Abisko 186 Quellgebiet 

Abisko 186 Quellgebiet 

Abisko 186 Quellgebiet 


Lappland 189 Abisko, Timpel 
Sumatra, SK Ic Singkaraksee 
Abisko 186 Quellgebiet 

Gr. Wasserfall, Moos. 7 
Neuseeland 

Lunz Nieder-Osterreich 
Tepl-Bett, Karlsbad 

Kolberg, Salzwiesen 2 

Abisko 157 3 Teich a Bahndamm 


Abisko 157 3 Teich a Bahndamm 
Schaalsee, Kirchenee Moos. | 
Wollingster Sea 1937 


Abisko, Lappland 157 Teich 

Domblitten 

Lunz Nieder-Osterreich 

Siskiyou County, California 

Auzillac Frankreich 

Gemindener Maar, Grund 

Pyrenaen: Lac d’Artouste Quell-sumpf 

Kitzbihel, Tirol 

Bei den Jffigenbachfallen Schweiz 

Spitzbergen Gletscher-Abfluss Moos 

Spitzbergen Gletscher-Abfluss Moos 

Java, DIC, Dieng Plateau, Telaga 
Merdodo 

Riesengebirge KI]. Schneegrube 

Abisko 186 Quellgebiet 

Abisko 157 3 Teich a Bahndamm 

Java, D 3a Dieng Plateau, Telaga 
Perigelon 

Sumatra SK Ic Singkaraksee 

Holzmaar an Ceratophyllum 

Pass Thurn, 6 an Dicranella an Felsen 
Tirol 

Pyrenaen Lac d’Artouste, Quell-sumpf 
18.b. 


Hustedt 
slide 
No. 
21 
54 
56 
14 
7), 

57 
Si 


58 
62 
57 
55 
59 
53 
25 
21 
66 


66 
9] 
24 


38 
29 
53 
94 
51 
52 
65 
32 
63 


342 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 13 


Denticula vanheurcki Brun 


Diploneis domblittensis (Grunow) Cleve 
Diploneis domblittensis var. 
subcontricta A. Cleve 
Diploneis elliptica (Kutzing) Cleve 
Diploneis marginestriata Hustedt 
Diploneis mauleri (Brun) Cleve 
Diploneis ovalis (Hilse) Cleve 
Diploneis pulcherrima Hustedt 
Epithemia sorex var. lapponica Hustedt 
Epithemia turgida (Ehrenberg) Kitzing 
Eunotia arcus Ehrenberg 
Eunotia bigibba Kitzing 
Eunotia clevei Grunow 
Eunotia denticulata (Brébisson) 
Rabenhorst 
Eunotia lapponica Grunow 
Eunotia serpentina Ehrenberg 
Eunotia triodon Ehrenberg 
Fragilaria capucina Desmazieres 
Fragilaria crotonensis (Edwards) 
Kitton 
Fragilaria javanica Hustedt 
Gomphocymbella ancyli (Cleve) 
Hustedt 
Gomphonema cantalicum Brun & 
Heribaud 
Gomphonema eriensee Grunow 
Gomphonema quadripunctatum 
(Ostrup) Wislauch 
Gomphonema subtile Ehrenberg 
Gomphonema transilvanicum Pantocsek 
Gomphonema ventricosum 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira arenaria Moore 
Melosira ruttneri Hustedt 
Navicula accommoda Hustedt 
Navicula accommoda Hustedt 
Navicula grimmei Krasske 
Navicula grimmei Krasske 
Navicula jakovljevici Hustedt 


Java, DIC Dieng Plateau, Telaga 
Merdodo 
Domblitten 
Nieder-Osterreich, Lunz Untersee 
Stat. I 
Nieder-Osterreich, Lunz Untersee V 11m 
Nieder-Osterreich, Lunz Untersee V 12m 
Domblitten 
Kitzbihel, Tirol 
Java TJ 2 IIc Tjibeureum Wasserfall 
Abisko 157. Teich 
Jmmerather Maar 1,5m 
Pyrenaen: Lac d’Artouste Quell-sumpf 
Eisenach, Drachenschlucht 
Setanai, Japan 
Abisko, Lappland, Tumpel 


Abisko, Lappland, Tiimpel 
Neuseeland 

Abisko, Lappland, Tumpel 
Wollin Plankton 

Behler See 


Sumatra, F.H. 2c Harau-Kloof Wasserfall 
Nieder Osterreich Lunz Untersee Stat. II 


Auzillac Frankreich 


Hainan 
Baikal See Jilischke 


Java D 3a, Dieng Plateau Telaga Pengilon 
Quelle bei St. Naum, Siidslavien 2.b 
Lunz Mittersee-Abfluss 

Quelle bei St. Naum, Siidslavien 
Eisenach 6233 

Tirol 3475 

Tirol 3480 

Tirol 3480 

Eisenach 6233 

Eisenach 6233 

St. Naum 735 

Wilhelmshohe bei Kassel 3790 
Eisenach 6232 

St. Naum 735 

Wilhelmshohe bei Kassel 3790 

Tirol 3475 

Java TJ2 IIc Tjibeureum Wasserfall 


Hemmentaler, Bach. Schaffhausen Schweiz 
Hemmentaler, Bach. Schaffhausen Schweiz 


Belg-Kongo. 529 May-ia Moto 
Kleinern bei Wildungen 
Quelle bei St. Naum, Stidslavien 2.b 


43 


29 
28 


22 
3a 
29 
32, 
Pa 
38 
41 
65 
37, 
40 
39 


MAHOOD: SOVEREIGN DIATOM COLLECTION 


Navicula ludloviana A. Schmidt 
Navicula perpusilla Grunow 
Navicula placentula Ehrenberg 
Navicula pseudobryophila Hustedt 
Navicula tenuicephala Hustedt 
Neidium distinctepunctatum Hustedt 
Neidium hitchcocki (Ehrenberg) Cleve 
Neidium meisteri Hustedt 
Peronia heribaudi Brun & Peragallo 
Pinnularia balfouriana Grunow 
Pinnularia gracillima Gregory 
Pinnularia polyonca (Brébisson) 
O. Muller 
Pinnularia pulchra Ostrup 
Pinnularia semicruciata (A. Schmidt) 
A. Cleve 
Pinnularia transversa (A. Schmidt) 
Synedra capitata Ehrenberg 
Stephanodiscus alpinus Hustedt & 
Ruttner 
Stephanodiscus astraea Ehrenberg 
Stephanodiscus carconensis Grunow 
Stephanodiscus damasi Hustedt 
Stephanodiscus dubius (Fricke) 
Hustedt 
Stephanodiscus hantzschia Grunow 
Stephanodiscus lucens Hustedt 
Stauroneis lauenburgiana Hustedt 
Stauroneis montana Krasske 
Stauroneis montana f. lanceolata 


Hustedt 
Stauroneis montana f. lanceolata 


Hustedt 
Stephanodiscus niagarae Ehrenberg 
Stephanodiscus niagarae Ehrenberg 
Stephanodiscus novaezeelandiae Cleve 
Stephanodiscus tenuis Hustedt 
Surirella amoena Pantocsek 


Surirella engleri O. Miller 

Surirella contorta Kitton 

Surirella obscura Reich 

Surirella skvortzowi Hustedt 

Surirella sovereigni Hustedt 

Surirella spinosa Hustedt 

Surirella spiralis Kiitzing 

Tabellaria binalis (Ehrenberg) Grunow 

Tabellaria fenestrata var. 
asterionelloides Grunow 

Tetracyclus rupestris (A.Brebisson) 
Grunow 


Fort Ludlow, Washington 


Pass Thurn 6 an Dicranella an Felsen, Tirol 


Nieder-Osterreich Lunz, Untersee V 11m 
Lappland 189 Abisko, Timpel 

Lappland 189 Abisko, Tumpel 
Nieder-Osterreich Lunz, Untersee V 11m 
Vatter-See 32 13m 17.8.26 Schweden 
Obere Kelle Gornergrat 

Riesengibirge Grosser Koppenteich 3557 


Pass Thurn 6 an Dicranella an Felsen Tirol 
Java, DIC Dieng Plateau, Telaga Merdodo 


Sumatra, FH 3D, Harau Kloof, Uberr. 
Wand 

Abisko 180 Schmelzwasser See 

Vatter-see 32 13m 17.8.26 Schweden 


Sodankyla a 
Jmmerather Maar 1, 5m 
R.S. Grundlsee Pl 5m 6.5 35 


Trentsee 

Pit River, Oregon 

Belg-Kongo 182 Edwardsee PI. 
Wollin Plankton 


Klinkerteich, Plon 

Ems. bei Hilkenborg 464 

Kuthrener Au Holstein 257 

Hasbruch Jagdhutte, Bach. Lebermoos 
Hasbruch, Wasserrinne, Moos 2 


Hasbruch, Wasserrinne, Moos 1 


Jsabol, Guatemala 

Eniesee, U.S.A: 

Kingsland, Neuseeland 

Ems, bei Papenburg 197 

Wehr bei Burgebrohl, Rheinland oberer 
Teil d. Lagers 

Belg-Kongo 134 Eduard-See Pl Obfl 

Taca Puna Neuseeland 

Puebla, Mexico 

Baikalsee 

Shadow Lake, King Co. Washington 

Java TJ 2 IIc, Tjibeureum Wasserfall 

Lunzer See 

Silbersee 10.8.36 

Viersee 26.8.17 


Bayrische Alpen Gfallermuhlen 4 Moos 
+600m 


a An irate’ Sah 
b« i“ A) 


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PROCEEDINGS 


OF THE 


Llonanite 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 14, pp. 345-356 


February 8, 1978 


NEW NORTH AMERICAN TABANIDAE (INSECTA: DIPTERA). XXIV. 
FURTHER COMMENTS ON CERTAIN PANGONIINAE IN 
MEXICO WITH SPECIAL REFERENCE TO ESENBECKIA 


By 


Cornelius B. Philip 


Research Associate, Department of Entomology, California Academy of Sciences, 
Golden Gate Park, San Francisco, California 94118 


ABSTRACT: 


Described as new from Mexico are: Esenbeckia (Ricardoa) leechi, female from Oaxaca, E. (R.) 


deltachi, female and male from Chihuahua, and Fidena flavipennis subspecies fisheri, females and male from 
Sinaloa to Guerrero. New taxonomic status includes new subspecies assignment of Esenbeckia melanopa 
(Hine) under E. semiflava (Wiedemann); and E. abata Philip, reassigned as a subspecies of EF. tepicana 
(Townsend). Significant new distributional data and a key to Esenbeckia species in Mexico are provided. 
Distributional and descriptive data are also given for Scione aurulans (Wiedemann). 


INTRODUCTION 


Taxonomic concepts are being progressively 
refined among the species in primitive, gener- 
alized groups assigned to the widely distributed 
subfamily Pangoniinae (Tabanidae). Certain New 
World groups, however, exhibit apparently re- 
cent radiation. Among these, the genus Esen- 
beckia is represented from the southern Nearctic 
through the Neotropical Regions. Intensified col- 
lecting in the northern part of its range has re- 
vealed active, obviously recent speciation, es- 
pecially in Mexico. Most species there, now 
assigned to the subgenus Ricardoa, have appar- 
ently diverged so recently that a few are difficult 
to assign, and the limits of the subgenus itself are 
in doubt. Nevertheless, it has taxonomic utility in 
the Mexican fauna, in spite of the rather arbitrary 
separation evidenced in the first couplet of the 
key appended below. 

At least some species have retained the postu- 
lated primitive habit of flower feeding. As a re- 


sult, collectors of flower-visiting flies, such as 
syrphids and bombyliids, as well as incidental 
insect-collecting botanists, have been prolific 
sources of material discussed in this and the pre- 
vious paper in this series (Philip 1973). Almost 
nothing else is known of the biology and biting 
propensities of these flies except for the southern 
United States reports which describe the peculiar 
(though unreared) larvae of two species that also 
occur in northern Mexico, viz., Esenbeckia in- 
cisuralis (Say) by Jones (1956) and E. delta (Hine) 
by Burger (1977). Jones also noted coincidental 
feeding on flowers by both sexes and on nearby 
cattle by females. 

My 1954 review included 24 species, sub- 
species, and varieties of Esenbeckia from 
Mexico, but the total has since been variously 
expanded to 33 plus the 2 species described be- 
low. A revised key is therefore provided to ac- 
commodate the additions. Most taxa presented as 
subspecies are only tentatively ranked because of 


[345] 


346 


insufficient available data on their ecological and 
geographical limits and degree of sympatry with 
other taxa. 

Species in two additional Mexican pangoniine 
genera are discussed. Distributional and de- 
scriptive data are given for Scione aurulans 
(Wiedemann) and a new subspecies is described 
for Fidena flavipennis (Krober). 


ACKNOWLEDGMENTS 


I am indebted to many persons and institutions 
for the loan of specimens, and to Drs. G. B. 
Fairchild and L. L. Pechuman for valuable in- 
formation. 

Specimens have been studied from the follow- 
ing institutional and private collections, and the 
abbreviations used in the text for these are: 
(CAS)—California Academy of Sciences, San 
Francisco; (CBP)—C. B. Philip, at California 
Academy of Sciences; (CNC)—Canadian Na- 
tional Collection, Ottawa; (DJP)—D. J. Pletsch, 
Mexico City; (EMF)—E. M. Fisher, at California 
State University, Long Beach; (GBF)—G. B. 
Fairchild, at University of Florida, Gainesville; 
(LLP)—L. L. Pechuman, at Cornell University, 
Ithaca; (NMNH)—National Museum of Natural 
History, Washington, D.C.; (UCB)—University 
of California, Berkeley; (UCD)—University of 
California, Davis. 


SYSTEMATIC ACCOUNT 
Genus Esenbeckia Rondani 


Esenbeckia (Ricardoa) leechi Philip, new species 


DIAGNosis.—A dark, robust species with sub- 
shiny, cherry-red, entirely black-haired abdo- 
men, orange antennae, bicolored red and black 
legs, and wings sharply dark brown in the ex- 
treme bases. 

DEsCRIPTION.—Holotype, female, length 14.5 
mm. Eyes bare. Frons tan pollinose, gently di- 
vergent below, index 1/3.0; a rather slender, me- 
dian, reddish keel reaching almost to anterior of 3 
ocelli at vertex. Subcallus flat, grayish pollinose. 
Face and cheeks dark brown, thinly dusted with 
gray pollen, and but little protuberant below an- 
tennae; beard mostly black with scattered, incon- 
spicuous pale hairs on lower cheeks. Scape and 
pedicel dark reddish, black haired; flagellum 
orange, the apical annulus elongated as usual. 
Palpus dark reddish, black haired with narrow 
lateral groove in middle third, short and pointed, 
just over a fourth length of proboscis. Latter a 
little longer than head height. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 14 


Thorax deep olive-brown, darker below, 
mostly black haired, with inconspicuous, scat- 
tered yellow hairs on notum, accentuated above 
wing bases; scutellum more reddish. Fore and 
hind femora and coxae black with concolorous 
vestiture; mid-femora more reddish; tibiae dark 
red, also with black hairs. Wings subhyaline, 
sharply dark brown basad of basal cells and hu- 
meral cross veins, costal cells faintly tinged out- 
wardly; cells R; closed and petiolate, M; wide 
open; the usual spur veins present. Halteres 
brownish red. 

Abdomen subshiny above and below, deep 
cherry-red, narrow median dark spots on first 
three tergites, entirely black haired without the 
usual pale terminal fringes or tufts. 

TyPe-SErRIES.—Holotype , male, Mexico, Oaxaca, Sierra de 


Miahuatlan, 2 km SE San José del Pacifico, 2438 m, 29-X-74 
(D. E. and J. A. Breedlove); CAS Ent. Type No. 12490. 


COMMENTS.—From other dark species of 
Ricardoa with wings sharply darkened in ex- 
treme bases, E. (R.) leechi is distinguished by its 
entirely black-haired, dark-reddish, subshiny ab- 
domen. There is plant pollen about the lower face 
and chest hairs of the holotype. The collectors, 
Dennis E. and James A. Breedlove are a father 
and son, the father, an Academy botanist (for 
whom another attractive, unique Ricardoa has 
been named), and the son, a young, active en- 
tomological collector. It is probable that the new 
species is at least a flower feeder like several 
other Ricardoa species. 

Named in honor of Hugh B. Leech, recently 
retired curator of Coleoptera in the Academy’s 
Department of Entomology. He has collected 
many interesting tabanids and has been a helpful 
professional friend over the years. 


Esenbeckia (Ricardoa) deltachi Philip, new 
species 

DIAGNOsIs.—A rather large, brownish-red 
species with considerable whitish vestiture; vari- 
able, inverted black mid-abdominal triangles; and 
bicolored legs. 

DESCRIPTION.—Holotype, female, length 17 
mm. Frons gray pollinose, nearly parallel-sided, 
index 1/3.6, a strong, thinly pollinose (partially 
worn), reddish, median carina nearly reaching to 
median ocellus. Subcallus, face, and cheeks gray 
pollinose; beard, pleura, chest, and fore coxae 
whitish pilose. Face moderately swollen to about 
level of mid-scape. Antennae bright orange, 


PHILIP: NEW NORTH AMERICAN TABANIDAE 


scapes a little longer than tall, with mixed dark 
and pale hairs. Palpi red with mixed black and 
some pale hairs, very short and pointed with 
broad flat bare lateral areas, about one-fourth 
length of proboscis. 

Unlined notum, scutellum and first abdominal 
segment dull brownish overlain with steel-gray 
pollen and scattered short pale hairs. Legs bicol- 
ored with mostly black hairs, femora black, 
tibiae brownish. Wings subhyaline, costal cells 
yellow, spur veins about twice as long as stems; 
cells R; characteristically closed and petiolate at 
margins. Halteres pale brown. 

Abdomen dull reddish brown with small, in- 
verted middorsal, dark, inverted triangles not 
crossing tergites 2 to 4; incisures pale haired, 
decreasing caudally to the entirely dark last two 
tergites; dorsum otherwise predominantly black 
haired. Venter reddish brown basally with mixed 
pale and black hairs grading to entirely black on 
last two sternites. 

Allotype, male, length 15.5 mm. In good 
agreement with holotype except for usual sex 
differences, and readily associated. Eyes bare 
with undifferentiated size of facets in contrast to 
the moderately enlarged area of upper facets in E. 
delta (Hine) males. Palpi more slender, and bare 
areas consequently reduced; about one-third 
length of proboscis, with mixed pale and black 
hairs. Black hairs on abdomen more extensive 
and pale-haired incisures more reduced. 


Type-SERIES.—From Mexico. Holotype, female, Chihua- 
hua, 4.8 km SE of Témoris, 25—VIIJ-69 (R. C. Gardner, E. S. 
Glaser, T. A. Sears) (UCD) and allotype, male, same locality 
and collectors as for holotype but 3.3 km N Témoris, 8-IX-69 
(UCD). 

Paratypes, 6 (2 males and 4 females), same Témoris locality 
and collectors: 1 female same data as holotype; 2 females, 
29-VIII-69; 1 female, 17—VIII-68, 1219 m; and 2 males, 3.2 km 
and 8 km N, 22-VIII-68 (UCD, CAS and CBP). 


COMMENTS.—The paratypes are from 15 to 17 
mm in length. Several have plant pollen about the 
bases of their mouth parts and on their faces. 
There is a tendency in some paratypes for the 
triangle on tergite 4 to widen into a dark basal 
band. The palpi in two females are more slender, 
as in the males; the palpi of one male are entirely 
black haired. Mid-frontal carina thinly pollinose 
in one female, more or less worn in the others. 

The possibility that this is another subspecies 
of E. incisuralis (Say) was at first considered, but 
this seems precluded by the white vestiture and 
other characters; also, the females do not have 


347 


the banded-appearing, more blackish abdomens 
of the smaller subspecies tinkhami Philip. The 
precinctive type-series of deltachi appears dis- 
tinguishable from related species by a combina- 
tion of larger average size, slightly narrower 
fronts in females, almost completely darkened 
femora, reduced pale hairs on abdomen, plus a 
row of inverted, dark triangles. E. micheneri 
Philip also has whitish vestiture anteriorly, but it 
is more extensively developed over the abdomen 
as well, two hind pairs of legs also pale haired, 
palpi about one-half lengths of proboscides, and 
fronts of females wider, their faces more swollen 
under the antennae. 


Comments on Other Esenbeckia Species 
Esenbeckia (Esenbeckia) divergens Philip 


This species was originally described (Philip 
1969) as a subspecies of E. filipalpis (Williston), 
the distribution of which is catalogued by Fair- 
child (1971) as Brazil, Paraguay, and Argentina. 
Since the only intermediate early record of Costa 
Rica has not been confirmed, my original sugges- 
tion is now followed that this is a distinct species 
from the west coast of Mexico, though the female 
is still unknown. 


Esenbeckia (Esenbeckia) minor Krober 


The type of this species from Costa Rica was a 
war casualty in Budapest. It had been in- 
adequately described as a variety of E. fascipen- 
nis (Macquart) from Brazil, but it is not so ac- 
cepted by present authorities. Until its identity 
can be confidently confirmed by other specimens 
from Costa Rica, a presumably similar species in 
Chiapas must remain of uncertain affinity; it is 
related to E. mejiai Fairchild of Central America, 
with similar broad, compact body, long palpi, and 
bare face, but it is more brownish with darker 
femora and halteres. 


Esenbeckia (Esenbeckia) wiedemanni (Bellardi) 


Twenty-nine females, much the longest series 
that I have seen of this sparsely represented 
species in collections, were taken recently in one 
day by Dr. Dennis E. Breedlove and son, James, 
during botanical studies on the southwestern 
slopes of Cerro Mozotal in Chiapas (23—XI-76, 
213 m, near Municipio Motozintla, CAS). They 
also netted four on the southwestern slopes of 
Vulcan Tekana (14—XII-76, 213 m, near neigh- 
boring Municipio Union, Chiapas). Additional 
single females were taken southeast of La 


348 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 14 


Trinitaria (2—XI—76, 1006 m) and from a foothill of 
Cerro Boquevan (15—XII—76, 2438-2743 m, also 
near Municipio Motozintla), deposited in CAS. 
The flies were captured mostly on flowers of 
Compositae without attempted attack on the col- 
lectors, and, curiously, without any males pres- 
ent at these blossoms. 

Bellardi thought that he was proposing a new 
name for a preoccupied Wiedemann species 
when he proposed the name wiedemanni. The 
two types in Turin, Italy, were redescribed by me 
in 1954. They have elongate, spatulate palpi. An 
additional specimen in the Turin collection has 
more pointed, Ricardoa-like palpi, and these 
palpi are also over half the length of the probos- 
cis. All of the present series have the pointed 
palpal variation and with partially sclerotized but 
flexible labella. The broadened, compact, sub- 
shiny abdomens make this another Mexican 
species intermediate with the subgenus Ricar- 
doa. 

Previous records from Guerrero and Chiapas 
may have been infrequent because the species is 
montane. A female collected by Dampf from 
Chemique, Chiapas, 15—XI-—34, is labelled in his 
writing *‘Jicqnojave’’ [?] ‘‘pic. batic,’’ which may 
refer to a mountain peak. A male from the previ- 
ously unlisted state of Oaxaca was also captured 
on a mountain and is herewith described as the 
neallotype of E. wiedemanni (Bellardi). 

DESCRIPTION OF NEALLOTYPE.—Male, length 
16.5 mm. Except for the usual sex differences, 
this agrees well with the female, and is readily 
associated. The eye facets are nearly homoge- 
neous in size. The palpi, though a little more slen- 
der than in the female, are rounded, not pointed 
apically, and about half the length of the probos- 
cis, which is also equal to about head height. The 
femora are obviously darker brown than the red- 
dish tibiae, with the wing fumosity likewise inten- 
sified baso-costally. The abdomen is subshiny, 
reddish brown, with pale hair tufts on outer cor- 
ners of tergites 2 to S. 

COLLECTION DATA FOR NEALLOTYPE.—Male, Mexico, 


Oaxaca, Cerro San Felipe, 1800 m, 18—X—48 (H. O. Wagner), 
CAS Ent. Type No. 13036. 


Esenbeckia (Ricardoa) semiflava subspecies 
melanopa (Hine) 


This species was based on two females from 
Sinaloa (no date). I have seen one other specimen 
from that state collected in October. Since de- 
scribing a melanistic variant of E. semiflava 


(Wiedemann) as var. melanista (from Michoacan 
and Jalisco, both collected in September), I have 
noted a similar divergence in color in E. 
melanopa, and likewise consider this also to be 
but a variant of typical E. semiflava, distin- 
guished by its yellow vestiture on cheeks and 
thorax. Of the strikingly bicolored, typical E. 
semiflava, | have seen three males and five 
females from Morelos, Michoacan, and Nayarit, 
all taken in October, and Fairchild (1971) adds 
Guerrero and Veracruz in his catalog. 

Material is still too meager to determine if there 
is any intergradation of forms, or any significance 
in the geographic separation, but to make the 
names available, they are here treated as sub- 
species. 


Esenbeckia (Ricardoa) semiflava subspecies 
melanista Philip 


The type female was taken in Michoacan. Re- 
cently two specimens of each sex, taken also in 
Michoacan, were supplied by Dr. W. W. 
Middlekauff; this enabled my describing the 
hitherto unknown male of the species. Two re- 
cently collected females agree with the holotype, 
as could be expected in this group; the two males 
show less melanism on the abdomen than in the 
female holotype. 

DESCRIPTION.—Neallotype, male, length 16.5 
mm. Eyes bare, holoptic with undifferentiated- 
sized facets. Head and thoracic characters as in 
the female, jet black with entirely black vestiture, 
the palpi likewise very short, about one-fifth the 
length of the proboscis. However, the abdomen is 
more yellow on tergites 2 and 3, plus the entire 
venter; a small inverted dark triangle nearly 
crosses tergite 3; tergite 4 is blackish with narrow 
yellow edges, as in the female; the black more 
restricted mesially on the following tergites. 

COLLECTION DATA FOR NEALLOTYPE.—Male, Mexico, 
Michoacan, ‘‘Cotija & Vic.,’’ 8-IX-—72 (E. A. Kane and B. 
Villegas), CAS Ent. Type No. 12865. 

COMMENTS.—The two males have extruded 
genitalia as though recently in copula. Plant pol- 
len was not observed on mouth parts of any of the 
four specimens. The second male differed from 
the neallotype only in having a narrow brown line 
in place of the median triangle on tergite 3. 

In addition to the entirely yellow abdomens 
with concolorous hairs behind the black tergite 1, 
the few males seen of typical semiflava have 
some scattered yellow hairs among the black 
ones on faces, chests, pleura, and fore coxae. 


PHILIP: NEW NORTH AMERICAN TABANIDAE 


Esenbeckia (Ricardoa) caustica (Osten Sacken) 


It is disappointing that, in spite of considerable 
recent collecting in Durango, no females have 
been seen of this large, mostly blackish species. If 
it shows the sexual dichromatism frequently 
found in other Ricardoa, the female would be 
expected to be at least 18 mm in length, probably 
black and black haired, including the beard, with 
reddish-orange antennae and tibiae, and perhaps 
reduced reddish spots on the sides of tergite 2. 
The species belongs in Ricardoa based on the 
proboscis (3.5 times longer than short black- 
haired palpi) of the type, which I examined in the 
British Museum in 1953. It should run in the key 
to the group with extreme bases of wings sharply 
dark. I have discussed (1954) the previous mis- 
taken synonymy. 


Esenbeckia (Ricardoa) schusteri Philip 


One female, additional to the type-series of a 
female and 2 males from Chihuahua, was recently 
taken by D. E. and J. A. Breedlove in Sinaloa, 3.2 
km S of Ocurahui, Sierra Surutato, 1981 m, 
1-X-70 (CAS). Pollen adherent to facial and 
chest hairs indicates probable capture on flowers. 


Esenbeckia (Ricardoa) tepicana (Townsend) 


Since my discussion (1954) of females but no 
recognized males, plus only the male and five 
female types of E. abata Philip, a considerable 
subsequent series of pertinent males and females 
now reveals that my original suspicion, that even- 
tually integradation might be discovered, was jus- 
tified, and that the status of E. abata should be 
reduced to that of a melanistic variant. The typi- 
cal, predominantly yellow-haired form occurred 
in three males, two females, and one intermediate 
from Morelos (the type state for E. abata; the 
allotype male, also mostly yellow haired though 
taken with the melanistic holotype and paratype 
females!), in two females and an intermediate 
female from Guerrero, in a female and an inter- 
mediate from Michoacan (September), and four 
males from Puebla (taken with two abata-form 
females). The last females were collected a month 
later in same locality where the only male, pre- 
sumed to be a melanistic variant in this dichro- 
matic species, had predominantly black vestiture 
in beard, on chest, and on dorsum of the darker 
reddish abdomen, plus dark brown halteres. An 
additional female of abata was dated November; 
except as otherwise noted, all the above were 


349 


taken in October between 1680 and 1980 meters 
in altitude, and most of both sexes carried flower 
pollen on head and chest hairs. 

The two forms thus fly together, but the ex- 
tremes in females are entirely different in appear- 
ance for key purposes and are treated as 
physiologic subspecies for taxonomic purposes. 


Esenbeckia (Ricardoa) planiventris (Macquart) 


After seeing and discussing (1954) only black- 
ish syntypes of this species in the Muséum Na- 
tional d’Histoire Naturelle, Paris, I have seen 
males from Sonora, Nayarit, Colima, Morelos, 
and a female from Guerrero. I have seen longer 
series of variety saussurei (Bellardi) (type from 
Veracruz seen in Turin), of either or both sexes, 
from Sonora, Nayarit, Colima, Jalisco, and Guer- 
rero. It is still too early to know if localized 
habitat influences account for variability, from 
the dark typical form with bicolored legs, to en- 
tirely reddish forms (including the legs), and to 
forms with various patterns of black to orange 
vestiture. 

Three large (17.5-19 mm) males, taken to- 
gether recently in Nayarit, 22.5 km NW of Tepic, 
elev. 550 m, 11—X—75 (Powell and associates) 
(UCB) emphasize this variation. One male is dark 
mahogany-brown with entirely brown to black 
vestiture and strongly bicolored legs; the second 
is more brightly reddish with orange hairs later- 
ally on all tergites, on the entire venter and beard, 
and the femora are a little darker reddish than the 
tibiae. The third is intermediate—brown abdo- 
men with lateral orange hairs only on outer cor- 
ners of tegites 4 caudad, beard brown, and femora 
darkened basally. Baso-costal infuscation is 
about the same in all. Such variants in sympatric 
individuals complicate their inclusion in keys, 
assuming they comprise a single species, 
planiventris; regardless, the name saussurei is 
useful for placing the different-appearing, lighter 
forms. 

This is another instance where the pre-1960 
concept of ‘‘variety’’ would have had utility for 
nomenclatorial and key purposes; for example, 
pending discovery of typical dark planiventris in 
Veracruz where the type of saussurei originated. 
While such incomplete information was _ ac- 
cumulating on these two ‘“‘species,’’ Fairchild 
originally believed them specifically distinct on 
palpal characters but catalogued (1971) them as 
subspecies, as I had considered them (1954), be- 
fore the extent of their distributional overlap was 


350 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 14 


known. In discussing variations in 11 specimens 
of saussurei from Guerrero, Williston (1901) 
noted, ** ... the abdomen varies in depth of 
colour, sometimes to almost black... .”’ 


Esenbeckia (Ricardoa) weemsi Phillip 


Until now, the holotype female from Sinaloa 
has remained unique. Six additional females were 
recently taken in Jalisco, 3.2 km N of Chapala, 
22—X-73 (S.C. Williams and C. L. Mullinex), and 
another from Puerto Los Mazos, 14.5 km N of 
Autlan, 29—VIII—70 (M. Wasbauer) (CAS). Some 
significant variation includes: length, 12-15 mm, 
dark trontal keels variably worn and prominent, 
hind tibiae not darker reddish than fore pairs, and 
abdominal colors vary from predominantly red- 
dish to black behind tergites 3 and entire venters. 
Plant pollen on the body of several specimens 
indicates flower visitation, but one collector re- 
ported the flies were netted about their persons. 


Significant New State Locality 
Records 


Esenbeckia (Esenbeckia): illota—Nuevo 
Léon, 2 males, dates not known; Campeche, 
female, May. 

Esenbeckia (Ricardoa): curtipalpis—Du- 
rango, male, female, Oct.; Tamaulipas, Jalisco, 
sexes not indicated (identified GBF); downsi— 
Sinaloa, female, Oct.; Nayarit, female, Sept.; 
flavohirta—Sinaloa, 2 males, female, no date: 
Jalisco, male, Oct.; Nayarit, male, Oct.; 
incerta—Jalisco, female, Oct.; Nayarit, 2 males, 
female, Sept., Oct.; Guerrero, male, Oct.; in- 
cisuralis var. tinkhami—Chihuahua, male, Aug.; 
nigronotata—Durango, male, female, Aug.; 
pavida—Mexico, Distrito Federal, female, Sept.; 
Michoacan, 3 males, female, Sept.; painteri— 
Sinaloa, female, Oct.; scionodes—Jalisco, 
female, Oct.; seminuda—Chihuahua, male, 4 
females, Aug., Sept. Detailed data for these 
specimens (which were sent to me by various 
individuals and institutions) are on file at CAS. 


Key to Mexican Esenbeckia 


la. Proboscides seldom either more than 
head height or more than twice as long as 
palpi; frequently rather slender bodied 
Ba Mis eR (Esenbeckia s. str.) 2 

Ib. Proboscides mostly some, or much 
longer than, head, and palpi seldom half 
their lengths; abomen usually robust, 


Das 


2b. 


San 


3b. 


4a. 


4b. 


Sas 


Sb: 


6a. 


6b. 


TA. 


Tho}. 


oftentinflated2ii2%/Laes5) eee 
Pheer fy fs (subgenus Ricardoa Enderlein) 8 
Abdomen predominantly unicolorous 
with varying patterns of pale pile...... 3 
Abdomen sharply bicolored, first 2 or 3 
tergites yellowish to red with varying- 
sized black macules mesially, black 
thereaftera. hes - tao. 2 eee 7 
Abdomen rather flat and broad, subshiny 
dark reddish to brown with lateral fringes 
or tufts of pale hairs; size about 15 mm 


Abdomen usually more elongated, paler 
and duller, yellowish to green in color; 
usually smaller, under 15 mm......... 5 
Wings sharply blackish, including squa- 
mae, basad of axillary incision; lateral 
hair tufts pale straw yellow to white; face 
thinly dull-gray pollinose (Guerrero) ... 
te wiedemanni (Bellardi) 
Infuscation expanded baso-costally, the 
squamae yellowish; outer tergal fringes 
brownish yellow; face ‘‘naked’’ (*‘Mex- 
IKOC MNES Pee ore fuscipes Enderlein 
Body and legs concolorous, pale yellow 
to brown 
Abdomen deep jade to paler green (or 
yellowish in faded specimens, at least 
hind femora and thorax darkened) (Brit- 
ish Honduras and Guatemala through 
Central “America;*)?Chiapas)} 922 5-eee 
LIEN CN ME SA) [ prasiniventris (Macquart)] 
Abdomen slender, pale brown, black- 
haired, except for pale-haired first seg- 
ment; legs entirely pale; palpi broadly 
sabre shaped (Nuevo Léon, Quintana 
Roo, Chiapas; Central America)....... 
ELISE SEA EAC AT illota (Williston) 
Abdomen mostly pale haired, propor- 
tionately unusually swollen, pale brown 
with vague pale incisural bands which 
widen caudally; coxae and femoral bases 
darker than thorax; palpi slender, 
pointed; (Durango; southern Texas) ... 
ts RAD TSE ee ee micheneri Philip 
Hind legs blackish; females with strong 
frontal keels (Guatemala; ?7Mexico).... 
FT Pe EW [translucens (Macquart)] 
Hind legs strongly bicolored; female fron- 
tal carina weak in upper front unless ac- 
centuated by wear (Nayarit) .......... 
FUR Ae ieee divergens Philip 


PHILIP: NEW NORTH AMERICAN TABANIDAE 


8b. 
9a. 


9b. 


10a. 


10b. 


lla. 


11b. 


12a. 


12b. 


13a. 


Sib: 


14a. 


14b. 


ISa. 


Sb 
16a. 


AALS SRE enspewa aes Sere a tances Bowls eeeterasare’ ef ake 43 
Cells in extreme base of wing (first M, 
first C) behind axillary incision, humeral 
cross vein in costal cell, and usually alula 
and squame as well, deep brown to black- 
ish, in sharp contrast with 2 basal cells 
(R, second M) and outer costal cell... 10 
Costal cell uniformly colored on both 
sides of humeral cross vein; if extreme 
base of wing darkened, this becomes dif- 
fuse outwardly 
Body sharply bicolored, blackish to and 
including abdominal segment 1, bright 
yellow from segment 2 caudad 
«cick ee ae ee semiflava (Wiedemann) 
Body not thus sharply bicolored, usually 
darkened, at least caudally .......... 11 
Dorsum of abdomen dark brown to 
blackish with tergite 2 in sharp contrast, 
bright yellow (rarely, only laterally) 
(Sonora, Michoacan, Sinaloa) 
4 Ue Ee aa (semiflava subspp.) 12 
Abdomen not with bright (or rarely dull) 
yellow girdle confined to tergite 2 .... 13 
Beard and thorax with vestiture strongly 
blackish; palp short and blunt, scape 
about as long as tall 
eae ee semiflava subsp. melanista Philip 
Thorax olive-brown with vestiture and 
beard yellowish; palp elongate, pointed, 
scape distinctly longer than tall 
ses semiflava subsp. melanopa (Hine) 
Body, appendages, and most vestiture 
dark brown to black, hind tarsi more red- 
dish; abdomen coarsely orange haired 
Canndailliy dee.aees sewers = ieee = Sake 14 
Not with this combination of dark body, 
blackish appendages, and dense orange 
Pile WeMiNG aoe oe tas Soe ee 15 
Dark brown to black, apical annuli of an- 
tennae reddish; orange pile extending 
forward to middle or more of tergite 2 
(Zacatecas to Guerrero) 
nets ns. Se ey pavida (Williston) 
Intensely black, including flagellums; 
caudal orange pile terminating abruptly 
on tergite 4 (Michoacan) .. painteri Philip 
Abdomen behind segment | dark ruby- 
red, entirely black haired, above and 
below (Oaxaca) leechi n.sp. 
Abdomen not colored as above ...... 16 
Large bee-like species with bright to 
pale-yellow pile on olive-drab to blackish 


16b. 


17a. 


17b. 


18a. 


18b. 


19a. 


19b. 


20a. 


20b. 


nN 
nN 
oy 


351 


thoracic integument, and pile thickset, 
blackish on abdomen; appendages red- 
dish brown, femora black (Durango) ... 
breedlovei Philip 
Not such large, blackish, yellow-haired 
SPECIES Gat AG. Sem erect oe 17 
Robust species, 15-18 mm; proboscides 
subequal to or little longer than head 
height, and palpi about % their lengths . 18 
Smaller, under 14 mm;_proboscides 
longer and palpi proportionately shorter 


apigsiliee SSN ARIES SNe RTM eS 20 
Frons broader than index of 1/3.0; face 
but little produced above, dull pollinose; 
hairs of palpi and antennal bases yellow; 
predominantly yellow-haired species 
(Durango) e7P Rane S gertschi Philip 
Frons narrower; face moderately pro- 
duced above to about level of mid-scapes 
and with subshiny areas; palpi and anten- 
nae black haired; more sparsely black- 
and-yellow-haired species with dark, 
unicolorous hind egs! 22ers 19 
Abdomen reddish on first 3 segments, 
mostly yellow haired with median, elon- 
gated dark spots (Sinaloa) . weemsi Philip 
Abdomen brownish black without pat- 
tern, mostly black haired basally, yellow 
hairs caudally (Chihuahua) 


schusteri Philip 
Venter and halteres yellowish red, body 
vestiture, often including palpi, predomi- 
nantly yellow (Nayarit to Morelos) (=? 
basilaris Wiedemann, 1830 not 1828).. 
tepicana (Townsend) 
Venter and halteres brownish black, 
body vestiture and usually palpi, black 
aireciet: HAE FETS Ba ero 21 


a. Beard and much thoracic vestiture pale 


yellow to creamy (Michoacan to Puebla) 
tepicana subsp. abata Philip 


_ Entire body vestiture blackish or only 


beard pale 
variant of tepicana subsp. abata Philip? 


_ Face entirely shining yellow; appendages 


and body mainly orange to honey-yellow; 
fronts subparallel-sided above with non- 
contrasting, reduced, bare median cari- 
NAC. Ae esc ME cet ols 23 


_ Face pollinose, often thinly so, at least 


under the antennae; if parafacials natu- 
rally shiny, body, appendages, and/or 


fronts not as above 24 


352 


23a. 


23b. 


24a. 


24b. 


Dias 


27b. 


28a. 


28b. 


29a. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 14 


Size over 13 mm; abdomen broad, trun- 
cate behind (occasionally with dark, in- 
definite, narrow median line on tergite 2 
and darkened caudally); frontal carina 
variably prominent in upper two-thirds 
according to wear; face protuberant to 
end of scape; palpi nearly one-half length 
of proboscis (Honduras; Chiapas?) .... 
fe paatees eae Mi Sarat seed yas [mejiai Fairchild] 
Size under 13 mm; abdomen more slen- 
der, rounded behind (without integumen- 
tal markings); frontal carina strong in 
upper three-fourths; face protuberant to 
about level of mid-scape; palpi shorter, 
about one-third length of proboscis (Oax- 
Aca) dees sat ose leon salt mexicana Philip 
Abdominal integument practically uni- 
colorous yellowish to black, vestiture 
frequently pale or orange............ 25 
Abdomen with some integumental pat- 
tern, at least dull red sides anteriorly, 
sometimes sharply bicolored or (at least 
first segment contrasting, pallid) with 
pattern ofimaculles =: Gee. asscesl cl... 33 


. Cells R; of wings narrowed but not closed 


(body including scutellum predominantly 
black, legs bicolored) (‘‘Mexiko’’)..... 
GEIS COM Bll H/o nto od ? fidenodes (Enderlein) 


. Cells R; characteristically closed ..... 26 
. Legs bicolored, or at least hind femora 


basally, broadly, and plainly darker than 
MDI ey hg AHA reed PD sradin LAly | D7 


- Legs uniformly colored, reddish or yel- 


low oniblackhained4t.. 80. SA yn) 32 
Medium-sized (14 mm), black-bodied, 
beelike species; femora one-fourth to 
one-third reddish distally; first 2-3 ab- 
dominal segments red with orange hairs; 
antennae and palpi orange-reddish with 
black hairs; face short, blackish, but little 
swollen (Tamaulipas) nitens Philip 
Differing in more than one of these 
chihactonges..2 965 20. Bia eae 28 
Abdomen entirely dark haired or with 
orange fringes on outer tergal corners; all 
femora darkened; wings markedly paler 
behind or in:some: cells sev22.:50.0).- 29 
Abdomen with at least paler-haired 
fringes or edges; chiefly hind femora 
darkened; wings more evenly fumose .. 
Eee on eee CO Oh ere eee ae ee 31 
Palpi short and stubby, not pointed, al- 
most one-fifth length of proboscis; vein 


29: 


30a. 


30b. 


Silas 


31b. 


32b. 


33a. 


33b. 


34a. 


margins strongly fumose (no lateral 
orange fringes) (Sonora) .............. 
curtipalpis Philip 
Not with this combination, and discs of 
several cells not especially paler 30 
Abdomen dark brown-black with bright 
orange outer fringes caudally; legs bicol- 
ored (Sonora to Guerrero) 
RTL AS Ae Aes os ee planiventris (Macquart) 
Abdomen lighter, from dark reddish 
brown with bicolored legs to entirely red- 
dish including legs, with or without vari- 
able patterns of orange hairs laterally and 
ventrally (Sinaloa to Guerrero) ........ 
.... planiventris var. saussurei (Bellardi) 
Size robust, about 15 mm, abdomen 
broad and truncated caudally (if sides 
obscurely reddish, see also couplet 39); 
proboscis markedly longer than head 
height, labella porrect and pointed; basal 
annulus of flagellum taller than scape 
(femora mostly black haired) (Chiapas) 
arlsiaelaaa tthe etter hoguei Philip 
Smaller, 14 mm or less; abdomen more 
rounded behind, less broad; proboscis 
but little longer than head, labella more 
ovoid, flexible; basal annulus subequal to 
height of scape (western coast of Mex- 
ico) downsi Philip 


OS ORCY CRON Get Cy Cra) (Ce (ct ich Oko 


Wir Ory ood 4 46 


S) fe) im) me) «\ le le) ei (ee, 10 fe) ‘elielis: =) ele hejia) {e 


a. Large reddish-brown species, mostly 


black haired with orange fringes on sides 
of abdomen (Nayarit) 
variant of planiventris (Macquart)? 


feral ia twiyie! fe) 6) (@) (eh \et\ev lal wi tat oti(a 


ele eeuee 


Smaller, bright yellowish-orange species 
with concolorous vestiture............ 
Set eee flavohirta (Bellardi) 


Unusually small (11 mm) compact 
species with short palpi; dark abdomen 
appearing banded because of accen- 
tuated, pale-haired incisures; frons with 
strong reddish median keel nearly reach- 
ing anterior ocellus; face shining brown 
except for thin pollen under antennae 
(GUCTIERO) ne eee keelifera Philip 


Not such small species with strong keels 
and, mostly banediaces:.... ee eee 34 


Cell M; unusually narrowed at apex (ad- 
ventitiously closed!) as much as or more 
than base of Cell My; abdomen black, 
predominantly pale haired, sides of ter- 
gite 2 dull reddish (Jalisco, Puebla) .... 
scionodes Philip 


sige ieee. eles! abie ful imitelveilie, je" (o\ivl sel ol iaiie 


PHILIP: NEW NORTH AMERICAN TABANIDAE 


34b. 


35a. 


35b. 


36a. 
36b. 
Bae 


37D: 


38a. 


38b. 


39a. 


39b. 


40a. 


40b. 


4la. 


4lb. 


42a. 


If Cell M, infrequently, moderately nar- 
rowed, then body colors otherwise ... 35 
Beard, chest, and fore-coxal pile whitish; 
robust usually gray-brown species often 
with rather small head (southern Nearctic 
and/or northern Mexico)............-. 41 
Hairs of at least chest and fore coxae 
yellowish to brown; more yellowish- 
brown species with proportionately 
fameen MEAS (04> cyecol-, eettatersscetaseraile oy 
Robust species, 17 mm or more 
Smaller species, less than 16 mm..... 38 
Abdomen polished ‘“‘reddish yellow to 
black,’ first segment dull olive-gray, 
concolorous with notum and scutellum; 
legs blackish (Chihuahua) 
Hes APM corse 5, orate eee seminuda (Coquillett) 
Abdomen with tergite 2 yellow-red and 
with a prominent inverted black triangle, 
brownish red basally to blackish cau- 
dally; legs usually bicolored (Durango) 
Gah. Bitte: Lease nigronotata (Macquart) 
Size about 15 mm; abdomen subshiny 
blackish behind with prominent pale in- 
CISULES PIES Pine we xd nice Ske we INT 39 
Size 14 mm or less; abdomen predomi- 
nantly yellow or dull blackish from tergite 


36 


40) 
Front parallel-sided without bare carina; 
femora pale haired (southern U.S.; 
northern Mexico) incisuralis (Say) 
Front divergent below with partially de- 
nuded carina; femora mostly black 
haired hoguei Philip 
At least hind legs bicolored; upper face 
swollen to end of scape, sides subshiny 
(Morelos to Oaxaca) .. triangularis Philip 
Legs yellowish, unicolorous; face dull 
pollinose, less swollen (Veracruz to 
GueELErO) ae sho eDamio’s! incerta (Bellardi) 
Palpi bluntly rounded at about half length 
of rather short proboscis, rufous like the 
antennae; notum bluish gray with pair of 
abbreviated, admedian pale lines an- 
teriorly (Esenbeckia s. str.?, see couplet 
[micheneri Philip] 
Palpi one-fourth to one-third length of 
relatively longer proboscis, falcate and 
tapered to a blunt point, bright orange- 
yellow like antennae; notum otherwise, 
une Osttecs eer atte s stormants -lepaysees Src 42 
Notum light brown with pale-margined 
scutellum; squamae tinted like adjacent 


42b. 


43a. 


43b. 


44a. 


44b. 


45b. 


46a. 


46b. 
47a. 


47b. 


48a. 


48b. 
49a. 


353 


alula; legs reddish; abdomen dull tan with 
wide pale incisures, but no median dark 
triangles though there may be sugges- 
tions of narrow mid-tergal lines (Sonora, 
Chihuahua; southern U.S.).. delta (Hine) 
Notum and scutellum darker; femora 
black, tibiae brown; abdomen dull red- 
dish with narrower pale-haired incisures 
and row of middorsal blackish inverted 
triangles (Chihuahua) deltachi n.sp. 
Cells in extreme bases of wings behind 
axillary incision, dark brown to blackish, 
sharply contrasting with costal cell be- 
yond humeral cross vein and 2 basal cells 
outward 
At least costal cell uniformly darkened or 
subhyaline and any dark infuscation not 
sharply confined to extreme bases of 
WIDES fern. Me oe eee. Oe racatee or 52 
Body sharply bicolored, blackish an- 
teriorly and bright yellow caudad of first 
segment; antennae red 
semiflava (Wiedemann) 
Body not thus sharply bicolored, black 
with mostly yellow abdomen; if occa- 
sionally so, flagellums and at least a spot 
ontergite2 blackia>. seen sae 45 


_ Thorax and first abdominal segment dark 


brown with mostly yellow hair; legs uni- 
formly brownish black; abdomen caudad 
of tergite 2 reddish brown with yellow 
margins, tergite 2 contrasting bright yel- 
low with an indefinite dark marking me- 
sially (Sinaloa) 
semiflava subsp. melanopa (Hine) 
Body not dark with such a contrasting 
yellow girdle on tergite 2 .......--.-- 46 
Abdomen and hind legs deep brownish 
black, last 3 to 4 segments and usually 
beard pale yellow pilose (Chihuahua) .. 
Pe is ot re nO or schusteri Philip 
Not with these characters ...-.-.---- 47 
Body and appendages dark brownish 
black, abdomen from tergites 2 or 3 and 
the last 2 sternites orange-red with con- 
colorous coarse pile (Guerrero, Micho- 
ACAI) t-kioctraaeneh Sstce pavida (Williston) 
Body colors otherwise, at least antennae, 
often palpi and tibiae red 
Size rather small, usually under 14 
mm 49 
Robust species, 15 mm and over 
Abdomen predominantly yellowish red; 


48 


49b. 


Sila: 


51b. 


Sas 


55b. 


S6a. 


56b. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 14 


some or all femora widely reddish apical- 
Lyre ees Sa ee tepicana (Townsend) 
Abdomen typically subshiny black, sides 
sometimes with dark ruby-red reduced 
laterally or expanded over most of ter- 
1 tepicana subsp. abata Philip 


. Abdomen plain, subshiny reddish brown 


with pale hair-tufts on outer corners of 
tergites wiedemanni (Bellardi) 


. Abdomen with middorsal black triangles 


OlSPOLS cw te hes oe eet ee eee core a =| 
Predominantly yellow, hirsute species, 
occasionally melanistic; abdomen dull 
reddish brown including first segment, 
black from segment 4 caudad (Durango) 
Pe Ne OE oa eee s gertschi Philip 
Chiefly black-haired species, first abdom- 
inal segment blackish but not caudal 
Sepments (DUFANZO) 20,0 560 sale os oor: 
caustica (Osten Sacken) 


. Cells R; narrowed but not closed (body 


including scutellum predominantly 
black, legs bicolored) (‘‘Mexiko’’)..... 
Sees ait ite oe fidenodes (Enderlein) 


. Cells R; characteristically closed and 


petiolate at wing margins............ 53 


. Face shining yellow (abdomen _ horn- 


yellow, often with indefinite dark median 
line on tergite 2) (Honduras; ? Chi- 


ADAS esses cece ees [mejiai Fairchild] 
. Face at least thinly dull pollinose 
PEERS 1 An a Se A 54 


a. Abdomenessentially unicolorous reddish 


brownetoblack 4... s440 eee sec ae 5)5) 


. Abdomen with prominent inverted dark 


triangles basally or modestly pale banded 
ONstercites 2 tora a8. Oe See. 57 
Legs blackish; 2 basal cells paler than 
costal cells; palpi one-third length of 
proboscides (Chihuahua) ............. 
seminuda (Coquillett) 
Legs red or strongly bicolored; 2 basal 
cells as dark as costal cells; palpi one- 
fourth or less length of proboscides ... 56 
Abdomen entirely blackish and black 
haired; scutellum blackish ......:..... 
Lee ee planiventris (Macquart) 
Abdomen brownish to bright red with 
orange-reddish hairs laterally, or these 
reduced or wanting; scutellum red with 
paler apex. c42' staan eee 
...planiventris subsp. saussurei (Bellardi) 


a. Large species, over 16 mm; abdomen be- 


57D: 


S8a. 


59b. 


60a. 


60b. 


6la. 


61b. 


hind second segment reddish brown to 
blackas ieee nigronotata (Macquart) 
Smaller, under 16 mm with more yel- 
lowish abdomen*.202:5 22 eee 58 
Abdomen pale brownish grading into 
broad pale bands across incisures (So- 
nora, Chihuahua; southern U.S.) 
delta (Hine) 


. Abdomen more yellowish red with dark 


basal trrangles/..-).:.- 20 ye eee 59 


. Upper face swollen to level of end of 


elongate scape, which is half again longer 
than tall, mostly black haired as are the 
palpi; latter not over one-fourth length of 
proboscis; entire costal and first M cells 
equally deep brown (Morelos to Oaxaca) 
triangularis Philip 
Face dull pollinose, but little swollen; 
scapes and palpi often yellow haired, the 
former subequal in length and height; 
palpi about one-third length of probos- 
cides; costal and basal cells paler yel- 
LOW, REO it, eee 60 
Legs uniformly yellowish; inverted 
triangle on tergite 2 narrower than tall and 
last 3 segments black with pale hair 
fringes; size under 13 mm (cell M; wide 
open) (Veracruz to Guerrero) 
UE EE. ASI ANOE. ime Fons incerta (Bellardi) 
Femora darkened at least basally, if pre- 
dominantly reddish, size larger and ab- 
domen not darkened caudally; middorsal 
triangle wider than tall .............. 61 
Size robust, 15 mm or more; abdomen 
frequently, and hind tibiae entirely, 
orange-red; cells M3 wide open (northern 
Mexico; southern U.S.) 


Ps ONC COR ONIRO COLON Och OG Osten 0 eG 


Oe sce Ooo 


© ee © «0 0 © ¢ ev 6, 0 


Sahih aoc 7 ea OIA ARES incisuralis (Say) 
Size smaller, usually under 14 mm; abdo- 
men usually blackish caudally; cells M3 
often narrowed at margin, as much or 
more than at base of Mz (adventitiously 
closed) Walisco, Puebla) @ 2) - ase 


Genus Scione Walker 


Scione aurulans (Wiedemann) 


Most records of this species from Mexico are 
for the three southernmost states. It also occurs 
in neighboring British Honduras and Guatemala 
(from whose Veracruz Peninsula came the types 
of the synonym Diclisa misera Osten Sacken). I 
have seen in various collections: 14 females from 


PHILIP: NEW NORTH AMERICAN TABANIDAE 


Chiapas, collected from 3 March to 29 April, and 
at altitudes (when listed) of up to 183 m; 3 from 
Oaxaca, | March and 29 May at 1219 m; and 3 
from Tabasco, | March and 3 April. In reducing 
S. lurida Enderlein from Colombia to a sub- 
species of S. aurulans, Fairchild (1966) called 
attention to its disjunct distribution, without con- 
tinuous intervening Central American records. 
Two females from Veracruz, Mexico, also 
suggest a variant similarly separated geograph- 
ically and perhaps seasonally, but these females 
are not described as a subspecies pending accum- 
ulation of more material. 

These are larger (15 mm) and darker than typi- 
cal brighter-yellowish aurulans, and the abdo- 
men is predominantly brown with black hairs and 
midrows of small, golden-haired triangles. Palpi 
are more slender and sticklike than the usual flat- 
tened, blade-shaped ones of typical aurulans. 
More specimens are needed from the area for 
assessment in this difficult scionine group. 


Genus Fidena Walker 


Fidena flavipennis Krober 
Fidena flavipennis fisheri Philip, new subspecies 


DIAGNosis.—A dark reddish-brown to nearly 
black, compact species with yellowish-brown 
wings, mostly black vestiture, including heavy 
beards except for bright orange to yellow or occa- 
sionally creamy-white hair tufts on outer edges of 
abdominal segments 5 and 6, and rarely on 2, 
above and below. Seldom some white hairs in 
front of wing bases or as fringes on upper calyp- 
ters in different combinations. Proboscides about 
as long as head and thorax together. 

DESCRIPTION.—Holotype, female, 18 mm. 
Eyes densely short pale pilose. Frons brown pol- 
linose, rather narrow, index 1/5.3, nearly 
parallel-sided, the usual triocellar tubercle at ver- 
tex. Frontoclypeus brown, sparsely grayish pol- 
linose with bare areas mesially, produced to 
about level of end of antennae. Two basal anten- 
nal segments dark brown, sparsely black haired, 
flagellums dull reddish, evenly tapered to the 
pointed apical annuli. Palpi brown, black haired, 
rather slender and pointed. Proboscis long and 
slender, black, peculiarly reddish on the porrect 
sclerotized labellums. 

Thorax brown, unpatterned, scutellum with 
reddish hind margin; pleura entirely black haired. 
Legs brown, black haired, tibiae a little more 
reddish. Wings smoky, more yellowish mesocos- 


355 


tally, the veins pale brown; cells R; closed and 
petiolate. No spur veins. Halteres brownish. 

Abdomen dark brown, polished dorsally with 
sparse black hairs, duller with more black hairs 
below; tufts of conspicuous coarse orange hairs 
on outer corners of tergites 5 and 6, extending 
farther inward on respective sternal incisures, a 
few unusual orange hairs on outer corners of ter- 
gite 2. 

Allotype, male, 19 mm. In good agreement 
with holotype but holoptic, the upper facets not 
enlarged; the partially extruded genitalia also 
with conspicuous orange hairs. It also lacks white 
tufts at base of wings, but has no yellow hairs on 
outer corners of tergite 2. 

Type-SERIES.—From Mexico. Holotype, female, Nayarit, 
vicinity of Higuera Blanca, about 9.5 km SW of Sayulita, 29- 
VII-71 (E. M. Fisher); and allotype, male, Sinaloa, 8 km NW 
of Escuinapa, 25—VII-71 (E. M. Fisher); both CAS Ent. Type 
No. 12491. 

Paratypes (55 females). Nayarit: 3, same data as holotype; 2, 
Rincon de Los Guayabitos, 31—VII—-70 (E. M. Fisher); 14, San 
Blas, various dates in August 1964 (J. F. McAlpine and others 
of the Canadian Mexican Expedition). Colima: 1, Playa de 
Santoya, 12.8 km NW of Manzanillo, 27—VIII-70 (M. Was- 
bauer). Jalisco: 3, Puerto Vallarta, ‘*8545°’ [S—VIII-45 ?] 
(Dampf Collection), and 5, same locality, Hotel Office, 28- 
VIII-76 (D. J. Pletsch); 1, 8 km SW of Copala, 975 m, 3—VIII- 
60 (R. A. Scheibner). Guerrero: 1, Carrizal, no date (W. G. 
Downs); and 3, Ixtapa, in building, 22—-VII-75 (D. J. Pletsch). 
Oaxaca: 22, Temascal, various dates in October 1965 (D. H. 
Janzen and R. F. Smith); (CAS, CBF, CNC, DJP, EMF, LLP, 
NMNH, UCB, UCD). 


COMMENTS.—This subspecies is_ cordially 
named for Mr. Eric M. Fisher in token of his 
supplying part of this type-series and many other 
Mexican tabanids. He supplied further collecting 
details on the allotype male: “collecting site is in 
area of nice Thorn Forest on gently sloping hill- 
side just above Hwy. 15; many Coleoptera were 
taken on flowering shrubs adjacent to mature 
forest; presumably the male was also taken on 
these flowers.’ If so, no pollen residue is now 
evident on mouthparts or heavy beard. 

The paratypes are in general good agreement 
with the holotype. In a few in which the stylets 
are unsheathed, the stylets nearly reach the bases 
of the peculiarly reddish, porrect labella. The 
caudal, outer-abdominal hair tufts occasionally 
may be pale yellow to creamy white, as in a 
topotypic female and five of the Temascal speci- 
mens; in only one of these, but in another 5 of the 
14 San Blas paratypes, there are some evident 
white prealar hairs, not seen in most of the others. 
Two of these have some yellow to creamy hairs 


356 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 14 


on the outer incisures of the second segments. 
Occasionally, the upper calypter is fringed with 
white hairs, not in any consistent combination of 
color characters. 

This new subspecies differs from the nominate 
Central American form, which substantiates geo- 
graphic separation, by the mostly bright orange to 
occasionally yellow or paler lateral hair tufts on 
the posterior abdomen, plus the usual lack of 
conspicuous white prealar tufts and average 
smaller size. Structurally, this subspecies seems 
to lie within some variation recorded for the 
nominate form by Fairchild (1942 and in litt.) 
farther to the south, but he also excluded a 
Nayarit specimen when he originally described 
F. isthmiae (=flavipennis). It was keyed and dis- 
cussed in my 1954 review as “‘isthmiae variant.’ 

The subspecies is distributed mainly along the 
Pacific Coast from Sinaloa to Guerrero, but the 
series from Temascal came from the Atlantic 
drainage in Oaxaca. Fisher reported taking the 
Higuero Blanco types around or on a decaying 
stream-side log at sea level, behind a mangrove 
swamp; the flies did not attempt to bite nor entera 
flight trap about 30 meters away. An altitude of at 
least 975 meters was recorded for the Copala, 
Sinaloa, specimen. 

Fairchild (1942) reported one of the paratypes 
of his isthmiae (=flavipennis) as captured while 


biting a boa constrictor in the Old Panama Zoo, 
and he recently (in litt.) reported specimens at- 
tempting to bite a recently killed, still-relaxed 
cayman. 

LITERATURE CITED 


BuRGER, J. F. 1977. The biosystematics of immature Arizona 
Tabanidae (Diptera). Trans. Am. Entomol. Soc. 103:145— 
258. 

Jones, C.M. 1956. Notes on the biology and feeding habits of 
Esenbeckia incisuralis (Say). J. Kans. Entomol. Soc. 29:43— 
46. 

FAIRCHILD, G. B. 1942. Notes on Tabanidae (Dipt.) from 
Panama. VIII. The genera Pityocera, Scione and Esenbeck- 
ia. Ann. Entomol. Soc. Am. 35(2):183-191. 

. 1966. Notes on Neotropical Tabanidae (Diptera) V. 

The species described by G. Enderlein. J. Med. Entomol. 

3:1-19. 

. 1969. Notes on Neotropical Tabanidae XII. Classifi- 

cation and distribution, with keys to genera and subgenera. 

Arq. Zool., Sao Paulo 17:199-255. 

. 1971. Family Tabanidae. In A catalogue of the Dip- 
tera of the Americas south of the United States. Mus. Zool., 
Univ. Sao Paulo, Brasil, fasc. 28. 163 pp. 

PuILip, C. B. 1954. New North American Tabanidae. VIII. 
Notes on and keys to the genera and species of Pangoniinae 
exclusive of Chrysops. Rev. Brasil. Entomol. 2:13-60. 

1969. New or little-known Neotropical Tabanidae 

(Diptera). Pan-Pac. Entomol. 45:147-152. 

. 1973. New North American Tabanidae (Diptera). XX. 
Six new species of Esenbeckia subgenus Ricardoa from 
Mexico. Ann. Entomol. Soc. Am. 66:1141-1145. 

WILLISTON, S. W. 1901. Supplement [concl.]. Pages 249-264 
in F. D. Godman and O. Salvin (editors), Biologia Centrali- 
Americana. Insecta. Diptera. Vol. 1. 378 pp., 6 pls. 


Vol. XLI, No. 15, pp. 357-370; 5 figs., 2 tables. 


PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 
FOURTH SERIES 


STRATIGRAPHY OF THE PLIO-PLEISTOCENE STRATA IN THE 


TWELVEMILE CREEK AREA, SAN FRANCISCO 
PENINSULA, CALIFORNIA 


By 
Thomas E. Yancey ay’ 


Department of Geology, Idaho State University, 
Pocatello, Idaho 83209 


ABSTRACT: Rocks in the Twelvemile Creek area are mostly sedimentary, belonging to the Franciscan, 
Merced, and Colma formations. The Merced Formation consists of about 1000 meters of richly fossiliferous 
marine strata, which correlate with the upper marine part of the type section of the Merced, and are entirely 
of upper Pliocene age. The nonmarine Colma Formation is subdivided into a lower massive sand unit a few 
hundred meters thick, and a thin clay-rich upper unit, and these units are suggested to have been deposited 
respectively, during a high stand and a low stand of sea level during the Pleistocene. The Merced and Colma 
formations in this area are separated by a high-angle fault, which is a northward continuation of the Serra 
Fault. The Serra Fault in this area nearly parallels the San Andreas Fault, gradually diverging from it towards 
the north. 

The Merced Formation in the Twelvemile Creek area is abundantly fossiliferous and contains a known 
fauna of over 50 species. The echinoids (Scutellaster), nassariid gastropods (Nassarius), and turrid gastro- 
pods (Ophiodermella) are the most useful fossils for correlation of Merced Formation strata. Species of the 
gastropod Ophiodermella have restricted stratigraphic ranges in the Merced Formation and probably can be 
used for regional correlations of Plio-Pleistocene strata. The species Ophiodermella graciosana (Arnold, 
1907), O. mercedensis (Martin, 1914), and O. incisa (Carpenter, 1864) occur in stratigraphic succession, and 
differ in having progressively weaker ornament and increasing spire height from O. graciosana to O. incisa. 


June 22, 1978 


INTRODUCTION 


The Twelvemile Creek area lies along the to- 
pographic crest of the San Francisco Peninsula 
and is characterized by high topographic relief 
with deep valleys incised into soft and semicon- 
solidated rock units, which have been uplifted 
along faults running in a NW-SE direction. The 
most important fault is the predominantly strike- 
slip San Andreas Fault, which has been inter- 
mittently active over a long period of geologic 
time and has had a controlling influence on de- 
position of late Cenozoic sediments in this area. 
The fault is believed to have controlled the for- 


mation of the depositional basin, its size and 
form, and to have influenced the types of sedi- 
ments deposited within it. 

The San Francisco Peninsula basin is one of 
several Pliocene or Plio-Pleistocene basins that 
formed along the northern California coastline 
during the time preceding the formation of San 
Francisco Bay. In early Pliocene times, subsi- 
dence along the San Andreas Fault created the 
depositional basin which now contains the 
Merced Formation on the San Francisco Pen- 
insula. A thin layer of nonmarine sediments was 
deposited unconformably over bedrock of Fran- 


[357] 


358 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 15 


SAN 
FRANCISCO 


\ 
a 


——, _ ic 
Hic TWELVE MILE crete saa) 


ANDREAS 


SG 
“RA FREEWAY 


Qls Landslide — Holocene ¥-—4- Fold 


° 
(._ _}}]0R Ravine fill — Holocene Quaternary 35 y¥ Bedding attitude 
° 
= Qc Colma Fm. — Pleistocene 90 Vertical bed 
(Es) Tm Merced Formation i; Tertiory (Pliocene) ® Horizontal bed 
ait Franciscon Formation i Cretaceous ONG 
—T~ — Foult showing dip (dashed where approximate) pees Stream 


FiGuRE |. Geologic map of the Twelvemile Creek area, San Mateo County, San Francisco, California. 


YANCEY: PLIO-PLEISTOCENE STRATA IN TWELVEMILE CREEK AREA 359 


ciscan Formation greenstones, followed con- 
formably by a change to marine deposition, with 
marine deposits accumulating continuously until 
the early Pleistocene when deposition became 
dominantly nonmarine and intermittent. 
Throughout this period, deposition took place in 
shallow-water conditions, and most deposition 
was very close to sea level. 

The late Cenozoic marine and nonmarine sed- 
imentary rocks in this basin, named the Merced 
Formation by Lawson (1893), are exposed in a 
long NW-SE trend parallel to the San Andreas 
Fault. The type exposures of the formation 
along the ocean cliffs to the northwest have been 
studied by Glen (1959) and Hall (1965a, 1965b), 
but there has never been any detailed work on 
the strata in the Twelvemile Creek area. The 
latter exposures provide the only stratigraphic 
section of the Merced Formation on the San 
Francisco Peninsula other than the type expo- 
sures along the ocean cliffs. 

As part of a study of the strata within this 
basin, a geologic map was prepared of the area 
around the head of Twelvemile Creek on the San 
Francisco Peninsula (Fig. 1). This area includes 
the San Andreas Rift Zone and the young sedi- 
ments northeast of the fault, encompassing an 
area between Skyline Boulevard on the west and 
Junipero Serra Boulevard on the east. The map 
is solely the work of the author except for some 
details of the trace of the San Andreas Fault 
taken from the geologic map of the San Fran- 
cisco South quadrangle by Bonilla (1965, 1971). 

This investigation began in the winter months 
of 1969 and continued through 1970, during a 
period of time when excavations combined with 
winter rains created favorable exposures over 
large areas within the Serramonte and Westbor- 
ough development districts in the towns of 
South San Francisco, San Bruno, and Pacifica. 
I am indebeted to Warren O. Addicott of the 
U.S. Geological Survey for encouragement and 
help in preparing the manuscript, and I thank 
him and the U.S. Geological Survey for photo- 
graphs of the fossils illustrated here. Also, col- 
leagues Maurice E. Kaasa and John Marr ex- 
pressed much interest and accompanied me in 
the field. 

Fossil locality descriptions are recorded in the 
Museum of Paleontology, University of Califor- 
nia, Berkeley. Illustrated fossils are stored in the 
U.S. National Museum and have received type 
numbers from their catalogs. 


MAJOR GEOLOGIC STRUCTURES 


Two major faults, the San Andreas Fault and 
the Serra Fault, control the structural trends in 
the Twelvemile Creek area. Both faults are ori- 
ented NW-SE, and the Serra Fault is probably 
generated from the San Andreas Fault, although 
both appear to be high-angle faults with signifi- 
cant vertical displacement. These faults cut the 
map area into three distinct structural blocks, 
each containing the exposures of a different sed- 
imentary formation. The structural block west 
of the San Andreas Fault contains metasedi- 
ments and greenstones of the Franciscan For- 
mation. The central block, between the San An- 
dreas and Serra faults, contains exposures of the 
Merced Formation which have been uplifted and 
tilted so that they now have a strike NW-SE, 
parallel to the San Andreas Fault, and dip steep- 
ly towards the north and east. The structural 
block east of the Serra Fault contains exposures 
of the Colma Formation which have not been 
significantly deformed and are essentially flat- 
lying and unconsolidated. 


San Andreas Fault 


The segment of the San Andreas Fault within 
the map area occurs as a well-defined rift zone 
of intensely sheared rocks a few tens of meters 
wide. Before construction work obliterated most 
of the topographic expression, the rift zone was 
identifiable by a series of sag ponds in the area. 
Exposures made during the course of construc- 
tion work showed that the actual limits of the 
rift correspond closely to the mapped limits as 
determined by topographic expression. 

The fault is tectonically active in this area 
(Bonilla 1959). The most recent significant activ- 
ity on the fault occurred during the 1957 San 
Francisco earthquake, although small earth 
tremors which are associated with the San An- 
dreas Fault system are a frequent occurrence. 
The last significant disruption of the ground sur- 
face along the fault occurred during the 1906 San 
Francisco earthquake when lateral displacement 
of a couple of meters developed along the fault 
(Lawson and others 1908; Bonilla 1959). 


Serra Fault 


The Serra Fault was named by Bonilla (1965) 
from surface exposures of a high-angle fault lo- 
cated about one kilometer south of the Twelve- 
mile Creek area. Within the map area this is a 
high-angle fault tending NW-SE approximately 


360 


parallel to the San Andreas Fault. It separates 
steeply dipping Merced Formation sediments 
from undeformed flat-lying Colma Formation 
sediments. Bonilla (1965) mapped the fault in the 
type area as a reverse fault with rocks of the 
Franciscan Formation thrust over rocks of the 
Merced and Colma formations. In the Twelve- 
mile Creek area the fault is clearly high-angle, 
but not demonstrably reverse or normal in ori- 
entation. It has a nearly linear trend, diverging 
gradually from the San Andreas Fault toward 
the north, and is in line with the segment of the 
fault mapped by Bonilla (1965). It has been 
mapped for a distance of over four kilometers. 

The Serra Fault lies close to the trend of the 
‘‘Foothills Thrust’? fault, a postulated north- 
south high-angle thrust fault lying east of the San 
Andreas Fault (Willis 1938). Bonilla (1959) dis- 
cusses this proposed fault, but does not use the 
name within the San Francisco South quadran- 
gle. The Serra Fault appears to be a secondary 
fault of the type proposed by Willis (1938), an 
upthrust fault connected to the San Andreas 
Fault at depth, bounding a wedge-shaped struc- 
tural block. Upward movement of the block 
from lateral compression could create thrusted 
fault boundaries. The upthrown side of the Serra 
Fault in the map area matches with the upthrust 
movement on the segment to the south, and sug- 
gests that the fault is an upthrust throughout its 
mapped length. 


Folds 


There is one conspicuous NW-SE tending fold 
within the Twelvemile Creek area. This is de- 
veloped in strata of the Merced Formation ad- 
jacent to the Serra Fault north of the north 
branch of Twelvemile Creek. It is a structural 
terrace consisting of a narrow band of strata 
with low dips, within a sequence of more steeply 
dipping strata. 


STRATIGRAPHY 


Three major stratigraphic units are present 
within the Twlevemile Creek area. The oldest is 
the Franciscan Formation, a mixed unit of 
greenstones and metasediments, of Jurassic— 
Cretaceous age. The next younger unit is the 
Merced Formation, a richly fossiliferous marine 
unit of Pliocene and Pleistocene age which was 
deposited on an eroded surface of the Francis- 
can Formation. The youngest unit is the Colma 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 15 


Formation, a flat-lying nonmarine unit of Pleis- 
tocene age that was deposited adjacent to or 
upon the eroded edge of the Merced Formation 
Strata. 


Merced Formation 


The Merced Formation was named by Law- 
son (1893) for a sequence of predominantly ma- 
rine Pliocene and Pleistocene strata on the San 
Francisco Peninsula. He divided the formation 
into lower and upper subdivisions, with the 
steeply dipping marine strata in the lower divi- 
sion. The type section of the Merced Formation 
was selected along the ocean cliff exposures 
south of San Francisco, from Mussel Rock in 
San Mateo County northward to Fleishhacker 
Zoo, north of the San Francisco City and Coun- 
ty line. 

The base of the type section at Mussel Rock 
has variously been considered to be a deposi- 
tional contact (Lawson 1893; Glen 1959; Hall 
1965a) or a fault contact (Ashley 1895; Higgins 
1961). Based on my own observations, I think 
that Glen and Hall are correct, and that it is a 
depositional contact. The lowest strata are clay 
rich, containing plant fossils, and rapidly change 
upward into marine deposits. These sediments 
are similar to that of modern long linear embay- 
ments along the San Andreas Fault (Bolinas 
Bay, Tomales Bay, Bodega Bay) and is an ex- 
pected depositional sequence for rising sea level 
transgressing into a long linear valley. The 
Merced Formation was probably deposited in a 
long valley within the San Andreas Rift Zone, 
since it is expected that linear embayments were 
present in Merced Formation time when the 
fault was also active. The upper contact of the 
Merced Formation was placed within a se- 
quence of deformed sandstones by Lawson 
(1893), while Hall (1965a, 1965b) included all of 
the deformed strata within the Merced Forma- 
tion and restricted the overlying unit (Colma 
Formation) to a thin layer of horizontal beds 
deposited on truncated Merced strata. The Col- 
ma Formation (of Schlocker and others 1958) 
consists of undeformed, poorly consolidated 
sandy deposits and is a partial equivalent of the 
‘*Terrace Formation’’ of Lawson (1895). 

Stratigraphic sections of the Merced Forma- 
tion in the Twelvemile Creek area are shown in 
Figure 3. The stratigraphic sequence extends 
from the San Andreas Fault on the west to the 
Serra Fault on the east, with only minor struc- 


YANCEY: PLIO-PLEISTOCENE STRATA IN TWELVEMILE CREEK AREA 361 


Kilometers 


FIGURE 2. 
Creek area. 


tural complications. The stratigraphic sequence 
is about 1000 meters (3000 feet) thick and is in- 
complete, with fault contacts at both the top and 
bottom. The lithology of the strata varies from 
clayey silts or clayey sands to shelly sandstones, 
with occasional thin peaty layers and thin con- 
glomeratic layers. The sediments are sparsely to 
richly fossiliferous and are normally poorly sort- 
ed. 

There is a trend of increasing average grain 
size of the sediments from the lower to upper 
part of the stratigraphic sequence. Sediments in 
the lower part of the sequence are mostly fine 
grained with some sandy interbeds, while sedi- 
ments in the upper part of the sequence are 
mostly sandy with occasional thin conglomeratic 


Stratigraphic 
A section 


D-3364 ®@ Fossil Locality 


Heavy mineral 
locality 


Location map of stratigraphic sections, fossil locations, and heavy mineral sample locations in the Twelvemile 


interbeds. The conglomeratic interbeds contain 
polished cherty pebbles and broken shell frag- 
ments and represent coarse-grained beach de- 
posits. Also, the upper, coarser sediments con- 
tain abundant echinoids, indicating deposition in 
high-energy conditions typical of beach environ- 
ments. This interpretation is further supported 
by the occurrence of many echinoid tests with 
the apical areas broken away, similar to the 
breakage of modern echinoid tests produced by 
waves on beaches, as can be seen on the nearby 
modern beaches of the San Francisco Peninsula. 

The Merced Formation in the Twelvemile 
Creek area contains both restricted and open 
marine biotas, characterized by the following 
species: 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 15 


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YANCEY: PLIO-PLEISTOCENE STRATA IN TWELVEMILE CREEK AREA 363 


Bivalves 
Mytilus condoni Dall, 1890 
Clinocardium meekianum (Gabb, 1866) 
Solen sicarius Gould, 1850 
Tresus nuttallii (Conrad, 1837) 
Macoma nasuta (Conrad, 1837) 
Gastropods 
Crepidula princeps Conrad, 1856 
Olivella biplicata (Sowerby, 1825) 
Polinices lewisii (Gould, 1847) 
Nucella lamellosa (Gmelin, 1790) 
Mitrella gouldii (Carpenter, 1857) 
Ophiodermella mercedensis (Martin, 1914) 
Crustaceans 
Cancer sp. 
Echinoids 
Scutellaster interlineatus (Stimpson, 1856) 
With the exception of Scutellaster, all species 
are living today or have closely related species 
living in the San Francisco area. The genus Scu- 
tellaster is extinct, but its ecologic niche appears 
to be presently filled by the genus Dendraster, 
as determined by a comparison of the modern 
occurrence of Dendraster and the inferred pa- 
leoenvironment of Scutellaster in the Merced 
Formation. 
Age.—An upper Pliocene age for this unit is es- 
tablished by the occurrence of Scutellaster in- 
terlineatus (Simpson, 1856), Nassarius morani- 
anus (Martin, 1914), and Mytilus condoni Dall, 
1890. The available evidence suggests that the 
genus Scutellaster became extinct by the end of 
the Pliocene (J. W. Durham, personal commu- 
nication, 1969) and is therefore an indicator for 
the Pliocene epoch. Nassarius moranianus is 
restricted to the upper Pliocene and lowermost 
Pleistocene (Addicott 1965). Mytilus condoni 1s 
restricted to the upper Pliocene and lower Pleis- 
tocene (Addicott 1974). Mandra (1949) reported 
it (as M. highoohiae Mandra, 1949) as a lower 
or middle Pliocene species, but its occurrence 
in the same horizon as Scutellaster interlineatus 
places it in the upper Pliocene (as used in central 
California). Nassarius moranianus has been 
found throughout the stratigraphic sequence, 
but Mytilus condoni and Scutellaster interlinea- 
tus have been found only in the upper part of 
the sequence. A bioseries of Ophiodermella is 
present in the Merced Formation, of which 
Ophiodermella mercedensis (Martin, 1914) is re- 
stricted to the Twelvemile Creek sequence and 
its correlatives, and is apparently restricted to 
the upper Pliocene. 


Correlation.—The Merced Formation strata in 
the Twelvemile Creek area correlate with the 
upper marine part of the Merced Formation in 
the ocean cliff section a few miles to the north- 
west (Figure 4). This is the second unit in Hall’s 
(1965a, 1965b) fourfold subdivision of the type 
section of the Merced Formation, which in- 
cludes strata lying between the San Andreas 
Fault and the marine-nonmarine transition in the 
formation. In both sections Scutellaster interli- 
neatus occurs near the top of the unit, and there 
is a trend of increasing grain size toward the top 
of the unit where coarse-grained beach deposits 
occur. Also, the same locally derived, heavy- 
mineral suite is present in the sands in both 
areas. The Twelvemile Creek section is about 
1000 meters (3000 feet) thick and is incomplete, 
compared to the same unit in the ocean cliff sec- 
tion which is about 1400 meters (4200 feet) thick 
and also incomplete. This is the thickest unit in 
the Merced Formation, and its outcrop can be 
traced continuously to the southeast from the 
ocean cliff section for about 13 kilometers (8 
miles) to the southern city limits of San Bruno 
and may extend discontinuously for several ki- 
lometers beyond that point. 

In the ocean cliff section the transition from 
older marine sediments to younger nonmarine 
sediments is marked by a change in heavy min- 
eralogy of the sands from a locally derived suite 
of sediments (basically from the Franciscan For- 
mation metasediments) to a distantly derived 
suite of sediments from the Sacramento-San 
Joaquin Valley (Hall 1965a). The Franciscan 
Formation-derived heavy mineralogy has a 
moderate proportion of hornblende and a very 
low proportion of hypersthene (Hall 1965a), di- 
agnostic Franciscan minerals such as jadeite, 
pumpellyite, lawsonite, and glaucophane in 
quantities up to 10-20 percent of the sample 
(Yancey and Lee 1972), and commonly has 
picotite and enstatite, also characteristic of 
Franciscan terrains. The Sacramento-San Joa- 
quin-derived heavy mineralogy has a lower pro- 
portion of hornblende and higher proportion of 
augite and hypersthene than the Franciscan-de- 
rived mineralogy. The proportion of augite plus 
hypersthene may reach SO percent of the sam- 
ple, with augite normally more abundant than 
hypersthene. Hall (1965a) demonstrated the 
source of the pyroxenes to be the late Cenozoic 
volcanics of the Central Valley and concluded 
that the sands of the nonmarine part of the 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 15 


TABLE 1. CHECKLIST OF MERCED FORMATION FossILs IN THE TWELVEMILE CREEK AREA. 


DO|— NIMS O/MW/O —|cum| +t w]}o 
DDI NOD NAD DVD DOWN Nanjyyn aja 
OO!O OH} OO/|/O O}|KM|DN DDD! MDM! HD 
Pole emelnelelelelnen 2) on ell el ell ol e)l'e) 
LO eI eT Ao 
aqgogogaqaqaoqaaoqoaqooo!)sa 


BIVALVIA 
Yoldia limatula 
Acila castrensis 


Mytilus condoni 
Mytilus coalingensis 


Mytilus sp 
Modiolus sp 


Adula californica 
Clinocardium meekianum 


Clinocardium sp 

Spisula albaria coosensis 
Spisula mercedensis 
Spisula mossbeachensis 


Tresus nuttallii 
Tresus sp 
Solen sicarius 
Siliqua patula 
Macoma nasuta 
Macoma sp 


Macoma inquinata 
Protothaca tenerrima 


Saxidomus sp 
Transennella tantilla 
Compsomyax subdiaphana 
Cryptomya californica 
GASTROPODA 


Bittium sp 


Epitonium tinctum 


Epitonium sp 
Calyptraea filosa 
Crepidula aff. C. princeps 
Polinices lewisii 

Nucella lamellosa 
Nucella canaliculata 
Nucella emarginata 
Neptunea tabulata 


Amphissa reticulata 
Mitrella gouldii 
Searlesia sp? 
Cancellaria sp ? 
Nassarius moranianus 
Nassarius sp 

Nassarius mendicus 
Olivella biplicata 
Ophiodermella mercedensis 
Ophiodermella graciosana 
Ophiodermella sp 
Turbonilla sp 


ARTHROPODA 


Cancer sp 


Callianassa sp 


Balanus sp 


ECHINOIDA 


Scutellaster interlineatus 
Scutellaster sp 


BRYOZOA 


YANCEY: PLIO-PLEISTOCENE STRATA IN TWELVEMILE CREEK AREA 


900m 


2500 


600m 


UPPER MARINE UNIT 


MERCED FORMATION 


300m 


Lower MARINE UNIT 


UNCONFORMITY 0 


Ww 
foal 
n 


= 
=) 
Ee 
aad 
== 
jo 
Be 
UNCONFORMITY a 
i tJ 
5500) = 
oo 
oe 
ao 


UNCONFORMITY 


a 


= 
TUFF BED z 
UPPER ae 
GASTROPOD =z 
BED 5000 £& 
J~= 
= 
Ss 
ZONE OF 
HEAVY 
MINERAL 1500 m 
CHANGE 
4500 
4000: 
1200 m 


3500 


|. _____—_—— vprer MARINE unite 
MERCED FORMATION 


3000 


FicureE 4. Columnar section of the Merced Formation strata from the type section of the formation (compiled from Glen 


1959, and Hall 1965a, 1965b). 


Merced Formation were derived from Sacra- 
mento-San Joaquin drainage that had newly es- 
tablished an outlet through the San Francisco 
Bay area. 

In the Twelvemile Creek area the heavy min- 
eralogy of the Merced Formation is the same 
throughout the section (samples 1-11 in Table 
2) and is similar to the mineralogy of the marine 


part of the Merced Formation in the ocean cliff 
section. It contains a basically Franciscan-de- 
rived heavy mineral suite with conspicuous 
glaucophane in the samples. A few of the sam- 
ples in this suite contain small but significant 
amounts of sphene and apatite, occasionally ac- 
companied by high hornblende content: min- 
erals indicative of granitic sources (Yancey and 


366 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 15 


TABLE 2. HEAVY MINERALOGY OF SANDS FROM THE MERCED (COLUMNS I-11) AND COLMA (COLUMNS 12-16) FORMATIONS 
IN THE TWELVEMILE CREEK AREA. Percentages are based on nonmicaceous-nonopaque minerals, and are based on minerals 


of approximately the same hydraulic character. Other minerals are tabulated separately. 


Samples 1 2 3 4 5 6 


Nonmicaceous, nonopaque minerals 


Hornblende 19 12 24 40 55 23 
Oxyhornblende 1 2 
Glaucophane 4 v7] 5 6 2 5 
Tremolite-Actinolite 1 3 3 1 
Hypersthene 1 5 2 3 1 
Augite 8 8 9 9 4 12 
Jadeite 10 11 25 18 14 6 
Enstatite 2 1 1 2 
Epidote 10 8 5 3 4 7 
Zoisite 1 1 
Pumpellyite 1 2 1 ] 
Lawsonite 3 1 4 
Garnet 7 8 2 3 1 10 
Apatite 2 1 4 3 22 
Tourmaline 3 
Zircon 8 6 6 4 10 
Sphene 4 5 5 6 12 4 
Rutile 1 1 
Picotite 19 23 4 3 3 11 
% 100 100 100 100 100 += 100 
Opaque minerals 
Magnetite-IIlmenite 52 36 9 13 18 40 
Hematite-Goethite 9 7 3 6 4 7 
Leucoxene 11 9 7 2 4 1 
Pyrite 1 1 
Micaceous minerals 
Biotite 2 1 2 3 3 
Chlorite 2 
Others 
Unknowns 5 6 6 4 2 6 
Carbonate 3 1 
Alterites 196 395 90 109 70 170 


Lee 1972). Granitic rock outcrops west of the 
San Andreas Fault were probably the source of 
these minerals. 


Colma Formation 


Unconformably overlying the Merced For- 
mation strata is a widespread unit of flat-lying 
undeformed deposits that were deposited on ir- 
regular topography. These deposits were de- 
fined and named by Schlocker and others (1958) 
as the Colma Formation, with a type area near 
the town of Colma, about 3 kilometers (2 miles) 
east of the ocean cliff section of the Merced For- 
mation. In its type area (where no significant 
section is presently exposed), the Colma For- 
mation is separated from the Merced Formation 


Uf 8 9 10 11 12 13 14 15 16 
37 19 = 43 25 39 «390 442i GB Ts 46 
2 1 3 3 
8 7 3 3 15 9 15 10 9 3 
3 5 1 2 9 4 9 9 6 9 

1 4 4 9 3 1 6 1 1 
12 14 13 10 12 16 12 11 16 18 
15 8 3 5 5 10 2 6 5 2 
4 2 2 1 2 1 
6 10 15 4 6 1 4 5 3 
1 
1 
2 1 2 1 
3 6 2 l 2 2 1 1 2 4 
2 1 3 
1 1 1 
1 >) 9 0 1 2 4 2 2 4 
9 7 8 3 2 2 2 2 3 3 
1 1 1 1 
3 4 1 19 4 6 2 6 8 5 
100 100 100 100 100 100 100 100 100 += 100 
14 =20 IS 45 4 23) 23 202i ees 
4 11 2 1 19 9 7 1 
1 9 2 6 4 8 1 i 
1] SS 9 1 
3 6 4 3 2 4 1 1 1 
88 273 165 173 192 439 330 84 330 280 


strata by the Serra Fault. The formation is prob- 
ably a few hundred meters thick in this area, 
although the thickest sequence reported by Bon- 
illa (1959) was 75 feet (23 meters), and the thick- 
est sequence measured in this study is about 60 
meters (200 feet). 

The Colma Formation is partly an equivalent 
of the ‘‘Terrace Formation’’ of Lawson (1893). 
Lawson gave this name to strata overlying 
his Merced Formation because he believed 
them to be terrace deposits derived from the 
underlying Merced. Schlocker and others (1958) 
renamed and redefined the strata of this interval 
because deposits from at least two tectonic ep- 
isodes are included in it. Most of the ‘‘Terrace”’ 
deposits along the ocean cliffs are most properly 


YANCEY: PLIO-PLEISTOCENE STRATA IN TWELVEMILE CREEK AREA 


placed in the Merced Formation as redefined by 
Hall (1965a, 1965b). 

The Colma Formation consists mostly of un- 
lithified sand of brown to yellowish color, and 
this is commonly the only lithology present in 
outcrops. A few pebble horizons can be found 
in most large outcrops, but they do not charac- 
terize any particular part of the section. In one 
zone near the road level of the Junipero Serra 
Freeway in South San Francisco (near heavy 
mineral locality 12), cobbles of a whitish, cherty 
rock are found in the sands. Scattered fossil 
fragments are found in the Colma Formation, 
but these could be reworked from Merced For- 
mation strata, and no complete shelly fossils 
have been found. Bonilla (1959) more fully de- 
scribes the lithologies of this part of the Colma 
Formation. 


Upper Part.—An upper part of the Colma For- 
mation is here distinguished for a group of strata 
that are lithologically distinct from the typical 
Colma but appear to be more or less conform- 
able with the typical Colma. This set of strata 
is probably equivalent to what Bonilla (1959) 
called the ‘‘Older Alluvium”’ in areas a few ki- 
lometers to the north, and to deposits of clayey 
sand in the Twin Peaks area of the San Francis- 
co North quadrangle mapped as alluvium by 
Schlocker (1974). This unit is characterized by 
sediments that are usually very poorly sorted 
and have a high clay content. The most common 
lithologies are poorly sorted, clayey, silty sands 
and sandy silts or clays containing abundant 
plant fragments. Bonilla (1959) noted that these 
strata are often more resistant to erosion than 
the underlying sediments and that they produce 
steep erosion slopes. These sediments are dis- 
tributed topographically above the friable sands 
of the lower Colma Formation. 

The clay-rich character of these sediments, 
their high organic (plant) content, and their very 
poor sorting indicate that they may be fossil soil 
horizons. Within the clay-rich horizons there is 
evidence of local channeling and reworking of 
deposits. Near heavy mineral locality 15 a very 
local unconformity with about 20 feet of relief 
was noted, and the overlying deposits contain 
horizons with clasts of unconsolidated clayey 
sand which are derived from immediately un- 
derlying deposits. This unit is probably the 
source of fossil tree trunks found about one ki- 
lometer south of Colma (Martin 1916). 


367 


Age.—Fossils of the Colma Formation (fossil 
woods) are not useful for age determination, but 
the entire formation can be dated as post-lower 
Pleistocene by stratigraphic position, since it 
overlies Merced Formation strata which have 
been interpreted to be as young as the Pleisto- 
cene Irvingtonian stage (Hall 1965a, 1965b). The 
main part of the Colma Formation was probably 
deposited during a higher stand of sea level, 
while the finer, clay-rich deposits probably ac- 
cumulated during a lower stand of sea level dur- 
ing the late Pleistocene. 

The sandy portion of the Colma Formation 
appears to be mostly reworked dune sands, 
judging from the rather uniform fine to medium 
grain size, moderate sorting, and lack of fossils 
or sedimentary structures apart from horizontal 
or subhorizontal bedding. Schlocker (1974:71) 
also noted that the Colma Formation deposits 
have about the same median grain size as dune 
sands. Assuming that the ocean beach was the 
main source for dune sands, deposition of the 
Colma Formation sands during a Pleistocene 
high stand of sea level is most probable since 
the source would be close to the area of depo- 
sition. Similarly, deposition of the upper fine- 
grained clay-rich deposits is most likely during 
a low stand of sea level when little wind-blown 
sand would be carried to the San Francisco Pen- 
insula from the distant ocean beach, which 
would be about 50 kilometers to the west at that 
time. 


Heavy Mineralogy of the Sands.—Colma For- 
mation sands (samples 12-16 in Table 2) studied 
for comparison with the Merced Formation 
show that the Colma Formation in the Twelve- 
mile Creek area has a composite heavy miner- 
alogy and that the minerals are extensively al- 
tered, probably by weathering before and after 
deposition. The sediments are not purely Fran- 
ciscan-granitic in origin nor purely Central Val- 
ley in origin, and are probably a mixture of both 
types. Colma Formation sands in the San Fran- 
cisco North quadrangle (Schlocker 1974:pl. 2) 
have a heavy mineralogy similar to that of the 
upper portion of the type Merced Formation 
(Hall 1965a), which is derived mostly from a 
Central Valley source area. These differences in 
heavy mineralogy in different areas apparently 
show local influences on sedimentation, but the 
majority of Colma Formation sands are probably 
derived from Central Valley sources. 


368 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 


15 


YANCEY: PLIO-PLEISTOCENE STRATA IN TWELVEMILE CREEK AREA 369 


AGE DIAGNOSTIC FOSSILS OF THE 
MERCED FORMATION 


Several groups of fossils are potentially im- 
portant for correlations of Merced Formation 
strata, especially the mytilids, myids, mactrids, 
nassariids, turrids, and echinoids, all of which 
have many species in the upper Cenozoic and 
have been reasonably well studied. The echi- 
noids and the turrids appear to be the most use- 
ful groups, having species with short strati- 
graphic ranges and occurring commonly in the 
Merced Formation. 

Species of the echinoid genus Scutellaster are 
common in coarse-grained marine strata and 
have restricted stratigraphic ranges. Glen (1959) 
records the stratigraphic ranges of Scutellaster 
oregonensis (Clark) and S. interlineatus (Stimp- 
son) in the ocean cliff section of the Merced, 
and points out the close relationship of the two 
species. It is probable that S. interlineatus 
evolved from S. oregonensis and that they have 
mutually exclusive biozones, which allows pre- 
cise correlation of Pliocene strata containing 
these species. 

Species of the turrid genus Ophiodermella 
also have restricted stratigraphic ranges in the 
Merced Formation and may be part of an evo- 
lutionary lineage including the species O. gra- 
ciosana (Arnold, 1907), O. mercedensis (Martin, 
1914), and O. incisa (Carpenter, 1864). The 
stratigraphically lowest-ocurring species is O. 
graciosana of which only a single individual was 
found close to the base of the section (loc. 
UCMP D-3693) in the Twelvemile Creek area 
and was not seen in the available collections 
from the ocean cliff section. The species has also 


— 


FIGURE 5. 


been found in exposures of the Merced? For- 
mation in northern Santa Clara County (Addi- 
cott 1969) that correlate with the oldest part of 
the Merced Formation in the ocean cliff section. 

The species O. mercedensis occurs commonly 
throughout the Twelvemile Creek section, oc- 
curring with O. graciosana only near the base 
of the section, and occurring alone in the upper 
part of the section. In the ocean cliff section it 
also occurs commonly throughout the section, 
and all of the O. graciosana var. mercedensis 
reported by Glen (1959) appear to be typical 
mercedensis. O. mercedensis is here considered 
a valid species and not a synonym of O. gra- 
ciosana, based on its weaker ribbing and ten- 
dency to be higher spired than O. graciosana. It 
does not show much variation in sculpture with- 
in the Merced Formation, with the intensity of 
spiral ribbing and radial ribbing being quite uni- 
form. The height of spire is slightly variable, and 
a few individuals show a noticeable weak keel 
in the position of the sinus. Glen’s (1959:178) 
statement that mercedensis varies greatly in 
spire height and ornamentation is based on ex- 
amination of badly weathered specimens, and 
these variations are not representative of the 
Merced Formation Ophiodermella. 

The living species O. incisa appears in the 
stratigraphically higher Upper Gastropod Bed of 
the Merced Formation. These three species are 
biostratigraphically restricted in the Merced 
Formation, and perhaps can be used for local 
and regional correlations. 

O. graciosana can be distinguished from O. 
mercedensis by its stronger and less numerous 
spiral ribs, presence of a weak keel, and radial 


Characteristic and distinctive fossils of the Merced Formation in the Twelvemile Creek area. A. Macoma 


inquinata (Deshayes) (0.7) USNM 251798, loc. UCMP D-5924. External view of specimen with mismatched valves showing 
dentition and rounded posterior margin. B. Macoma nasuta (Conrad) (0.6) USNM 251799, loc. UCMP D-3688. C. Mytilus 
condoni Dall (0.7) USNM 251800, loc. UCMP D-3364. View of hinge of a large valve. D. Mytilus condoni Dall (0.6) UCMP 
10928, loc. UCMP D-5927. Specimen with paired valves showing strong divaricating sculpture, but partly distorted due to 
sediment compaction. E. Transennella tantilla (Gould) (5) USNM 251801, loc. UCMP D-3386. F. Transennella tantilla (Gould) 
(3.2) USNM 251802, loc. UCMP D-3386. Slightly oblique view of average size specimen. G. Spisula albaria coosensis Howe 
(0.6) USNM 251803, loc. USGS Cenozoic M-5754. Internal and external views of a valve showing hinge and growth lines. H. 
Cryptomya californica (Conrad) (0.7) USNM 251804, loc. UCMP D-3688. I. Crepidula aff. C. princeps Conrad (0.7) USNM 
251805, loc. UCMP D-3364. Oblique apical view of a typical specimen. J. Nucella canaliculata (Duclos) (0.7) USNM 251806, 
loc. UCMP D-S765. Side view of a very elongate specimen. K. Nassarius moranianus (Martin) (0.7) USNM 251807, loc. UCMP 
D-3688. L. Amphissa reticulata Dall (1.4) USNM 251808, loc. UCMP D-3690. M. Bittium sp. (1.4) USNM 251809, loc. UCMP 
D-5920. N. Bittium sp. (1.4) USNM 251810, loc. UCMP D-5920. Side view showing sculpture with secondary spiral ribs on 
body whorl. O. Ophiodermella graciosana (Arnold) (1.4) USNM 251811, loc. UCMP D-3693. Side view of fragmentary specimen 
showing sculpture of strong nodes on whorls. P. Ophiodermella mercedensis (Martin) (1.4) USNM 251812, loc. UCMP D-3364. 
Apertural and abapertural views of a typical specimen. Q. Ophiodermella incisa (Carpenter) (1.4) USNM 251813, loc. UCMP 
B-4811 (from the Upper Gastropod Bed, at Thorton Beach State Park). 


370 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 15 


ribs across the upper half of the whorl, and O. 
incisa can be readily distinguished from O. mer- 
cedensis by its higher spire and fine spiral ribs, 
and lack of radial ribs except on the tip of the 
spire. These species show trends of increasing 
spire height, increase in spiral ribs, decrease in 
strength of radial ribs, and decrease in inflation 
of whorls through geologic time. 


LITERATURE CITED 


ADDICOTT, WARREN O. 1965. Some western American Cen- 
ozoic gastropods of the genus Nassarius. U.S. Geol. Surv. 
Prof. Pap. 503-B: B1—-B21. 

1969. Late Pliocene mollusks from San Francisco 

Peninsula, California, and their paleogeographic signifi- 

cance. Calif. Acad. Sci., Proc., Ser. 4, 37(3): 57-93. 

1974. Recognition and distribution of Mytilus con- 
doni Dall, a unique Pliocene and Pleistocene bivalve from 
the Pacific coast. Veliger 16(4): 354-358. 

ASHLEY, GEORGE H. 1895. The Neocene stratigraphy of the 
Santa Cruz Mountains of California. Calif. Acad. Sci., 
Proc., Ser. 2, 5(1): 273-367. 

BONILLA, MANUEL G. 1959. Geologic observations in the 
epicentral area of the San Francisco earthquake of March 
22, 1957. In G. B. Oakeshott, ed., San Francisco earthquake 
of March, 1957. Calif. Div. Mines Geol., Spec. Rep. 57: 25- 
She 


. 1965. Geologic map of the San Francisco South quad- 

rangle, California. U.S. Geol. Surv., Open File Report. 

. 1971. Preliminary geologic map of the San Francisco 
South quadrangle and part of the Hunter’s Point quadran- 
gle, California. U.S. Geol. Surv., Miscellaneous Field Stud- 
ies Maps MF-311, 2 sheets. 

GLEN, WILLIAM. 1959. Pliocene and lower Pleistocene of the 


western part of the San Francisco Peninsula. Univ. Calif., 
Publ. Geol. Sci. 36(2): 147-198. 

HALL, N. TIMOTHY. 1965a. Petrology of the type Merced 
Group, San Francisco Peninsula, Calif. Unpubl. Master’s 
Thesis. Univ. Calif., Berkeley. 127 pp. 

. 1965b. Late Cenozoic stratigraphy between Mussel 
Rock and Fleishhacker Zoo, San Francisco Peninsula. Jn 
Guidebook for northern California, 7th INQUA Congress. 
Nebraska Acad. Sci.: 151-161. 

HIGGINS, CHARLES G. 1961. San Andreas Fault north of San 
Francisco, California. Geol. Soc. Am., Bull. 72: 51-68. 

Lawson, ANDREW C. 1893. The post-Pliocene diastrophism 
of the coast of southern California. Univ. Calif. Publ., Bull. 
Dep. Geol. 1(4): 115—160. 

1895. Sketch of the geology of the San Francisco 

Peninsula. U.S. Geol. Surv., Ann. Rep. 15: 405-476. 

, AND OTHERS. 1908. The California earthquake of 
April 18, 1906. Report of the State Earthquake Investigation 
Commission. Carnegie Inst. Wash., Publ. 87, 1(pt.1): 1-451. 

MANDRA, YORK T. 1949. A new species of Mytilus from the 
Pliocene of Humboldt County, Calfironia. J. Paleon. 23(1): 
104-105. 

MaRTIN, BRuCcE. 1916. The Pliocene of middle and northern 
California. Univ. Calif. Publ., Bull. Dep. Geol. 9(15): 215- 
259. 

SCHLOCKER, JULIUS. 1974. Geology of the San Francisco 
North quadrangle, California. U.S. Geol. Surv., Prof. Pap. 
782: 1-109. 

, MANUEL G. BONILLA, AND DorotHy H. RAp- 
BRUCH. 1958. Geology of the San Francisco North quad- 
rangle, California. U.S. Geol. Surv., Misc. Geol. Invest.: 
Map I-272, 1 map with text. 

WILLIS, BAILEY. 1938. San Andreas Rift, California. J. Geol. 
46(6): 793-827. 

YANCEY, THOMAS E., AND JAMES W. LEE. 1972. Major 
heavy mineral assemblages and heavy mineral provinces of 
the central California coast region. Geol. Soc. Am., Bull. 
83(7): 2099-2104. 


. oe 
ote whet ile aan oe ae ay 
@ e 7 me Da 6) a wie 7 i haeae a 
= 7 a oe ee aa h=« WO Fue ros 
fun; Je Push 
; fh Pe eS ere eu Ve set 
yesdby & @ se Gat = i 


; bd. on as % io hee Le 
x 7 a (a a wey i<est 
: olan ars.. diy Qijaly 


—_ - ' 4 fF & ae ¢s 
ed 
s ae A 
om, 5 4 Tis of @ - 
A 4 fs 
» Par, Fee vee 
a 42 aa 7 os te ab 
-_ i) > a ue | ° 
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; fom 7! a ie? re tox : a — oe Sictba ae 
ey (ota ie” SP : bal OL Vine, Aiereey eo 
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OWA E ly Big Lomi ; : ‘ i & Ww ie ¢ Gira 


ee ee A ee : =. r na @ Wl nt or : 
Pa A a oa al ; ) E 
ry (i 3s a meted ~ Canes ae ” 4 
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PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 16, pp. 371-387; 51 figs., 2 tables. 


December 28, 1978 


THE STOMATAL COMPLEX IN AGAVE: GROUPS DESERTICOLAE, 
CAMPANIFLORAE, UMBELLIFLORAE 


By 
Howard Scott Gentry & Jane R. Sauck* 
Desert Botanical Garden, Papago Park, Phoenix, AZ 85010 


ABSTRACT: 


The stomatal cell complexes of 23 taxa in the genus Agave are described, measured, and 


illustrated. The taxa belong to three taxonomic groups preponderant in Baja California and other parts of 
the Sonoran Desert region: Deserticolae, 17 taxa; Campaniflorae, 3 taxa; Umbelliflorae, 3 taxa. 

The stomatal characteristics are compared with the contemporary systematic revision of Gentry (1978) at 
both the sectional and species levels. Stomatal differences are significant and helpful to Agave taxonomy at 
the sectional level, but are found to be irregular as species criteria. One species, Agave fortiflora, on the 
basis of stomatal anatomy, was found misaligned with the Umbelliflorae and was removed to another section. 

The apparent xeromorphic structure of agave stomates are compared with rainfall regimes of respective 
species. No correlations are found between apparent xeromorphic stomatal structures and the most arid 


habitats of agave species. 


INTRODUCTION 


During the course of making a taxonomic re- 
vision of the genus Agave of the California Gulf 
region, Gentry (1978) made a classification 
based on the gross morphology of the leaf, the 
inflorescence, the flower at anthesis, and the 
growth and reproductive habits. It became clear 
that levels of taxa and their perimeters were sub- 
tle and hard to define. Therefore, a critical study 
of the epidermis was made to see if such char- 
acters could be of correlative value in the pro- 
posed system. 

Characters of the epidermis and stomata have 
been found to be of considerable value in iden- 
tification and classification in various flowering- 
plant groups by Stace (1966), Tomlinson (1974), 
and others. Various accounts of the anatomy of 
Agave leaves include descriptions of epidermis 
and stomata. Probably the most extensive is that 


* Present address of J. R. Sauck: Dept. of Veterinary Sci- 
ences, Rm 232 Ag. Sci. Bldg., University of Arizona, Tucson, 
AZ 85721. 


of Miller (1909) in which about 95 species of 
Agave are described and a key devised for their 
separation and identification. Since no study ex- 
ists for Agave species inhabiting the California 
Gulf region, we examined the twenty-three taxa 
which occur in that area. The taxa fall into three 
sections or groups of the genus Agave: the De- 
serticolae, the Campaniflorae, and the Umbel- 
liflorae. A Sonoran Desert species, A. fortiflora, 
is also examined as to its relationship with the 
Umbelliflorae with which it was placed in an ear- 
lier paper by Gentry (1972). 

More recently, attempts have been made to 
classify and characterize Agave stomata for 
phylogenetic purposes (Stebbins and Khush 
1961; Shah and Gopal 1969), but as yet only one 
other study successfully utilized leaf anatomy, 
including the study of epidermal characters, to 
distinguish genera in the Agavaceae (see Blun- 
den, Yi, and Jewers 1973). 

The Deserticolae includes 17 taxa which are 
found in the Sonoran Desert in southeastern 
California, Arizona, Baja California, and Sono- 


372 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 16 


FIGURE 1. Surface view of the stomatal complex of Agave 
sobria ssp. roseana (x400). P,, polar subsidiary cells; L, lateral 
subsidiary cells. 


ra. Most species have a slender inflorescence 
bearing small, compact umbels and are either 
suckering or simple multiannuals. The short, 
open flower tube is characteristic. Nearly all are 
xerophytic. The Campaniflorae, native to the 
southern half of peninsular Baja California, con- 
sists of three species with broad campanulate 
flowers borne in panicles with smaller bracts. 
The Umbelliflorae consists of three taxa found 
in Baja California. They are distinguished by 
flowers borne in large umbellate branches sub- 
tended by conspicuous sheathing bracts, and 
they are mainly large perennials branching from 
the leaf axils. 


MATERIAL AND METHODS 


Materials were obtained primarily from her- 
barium specimens, though a few specimens were 
obtained from living material at the Desert Bo- 
tanical Garden, from Gentry’s personal collec- 
tion in Murrieta, California, and one from Santa 
Barbara Botanic Garden, California. Fresh ma- 
terial was preserved in Craf’s III or formalin- 
acetic-ethanol. Although one slide was prepared 
for each species, numerous herbarium speci- 
mens were examined under the dissecting mi- 
croscope to insure that a representative sample 
was taken. In addition, leaf peelings of over 40 


FIGURE 2. 


Cross-section of epidermis of Agave deserti. A, 
margin; B, cross-section of the polar lip; C, longisection of 
the lateral lip; D, chamber below the lateral lips (inner aper- 
ture); EF, stoma. 


species representing most other sections of the 
genus were prepared for general comparative 
purposes. 

Peelings of leaf epidermis were obtained 
readily from either preserved or dried material. 
Dried material was boiled 20 minutes in a deter- 
gent solution to soften it and to facilitate the 
removal of underlying tissues. Such material 
was rinsed in distilled water prior to dehydration 
and staining. Permanent mounts were made 
from peelings dehydrated in ethanol and stained 
ina 1% solution of Safranin O and 0.5% solution 
of Fast Green and mounted in Harleco Synthetic 
Resin. 

An attempt was made to prepare plastic im- 
prints according to the method of Sinclair and 
Dunn (1961). Due to the relatively deep supra- 
stomatal chamber of most species examined, the 
method proved to be unsatisfactory, as has been 
noted for Agave by Shah and Gopal (1969). 
Though the imprint method is not suitable for 
detailed morphological studies, sufficient detail 
was obtained to count stomata. 

After examining the leaves from several spe- 
cies, a uniformity was noted with regard to the 
structure of the stomata. Therefore, a single leaf 
was used. Five stomata were measured for each 
leaf. These measurements were taken about 
one-half way up from the base of the leaf. 

Leaf sections were required in order to inter- 
pret surface features of the epidermal peelings. 
Such sections were prepared from material pre- 
served in Craf’s III or FAA and embedded in 
paraffin, and dehydrated and stained in 1% Saf- 
ranin O and 0.5% Fast Green according to the 
method of Johansen (1940). Sectioning was im- 


GENTRY & SAUCK: STOMATAL COMPLEX IN AGAVE 


proved in hardened material by soaking the cut 
block in cold water for one hour to several days. 


GENERAL MORPHOLOGY OF THE 
STOMATAL COMPLEX 


In Agave, four epidermal cells surround the 
guard cells. These cells appear different from 
other epidermal cells in that two of them lie po- 
lar and two lie lateral to the guard cells. Often 
they stain differently from other epidermal cells, 
and they may possess a distinctive bulbous lip 
which protrudes into a cavity above the stoma 
called the suprastomatal chamber. Morphologi- 
cally, the stomatal complex appears to be tetra- 
cytic, but as noted by Blunden et al. (1973), the 
term is purely descriptive as developmental 
studies have not been carried out in this paper. 
We will refer to these cells as subsidiary cells 
following Tomlinson (1974), who retains the 
term for structurally specialized cells distinct 
from other epidermal cells associated with the 
guard cells of the mature stomata (Fig. 1). The 
guard cells and associated subsidiary cells are 
oriented parallel to the long axis of the leaf. 

The stomatal-complex proper lies at the bot- 
tom of a cavity, the suprastomatal chamber, 
created by the various elaborations of supra-epi- 
dermal structures, which consist of a dark- 
staining waxy or cellulosic layer capped by a 
red-staining cuticular layer (Fig. 2). That the sto- 
mata are truly sunken is questionable; the sto- 
mata are actually depressed below the cuticular 
and subcuticular layers only and just slightly 
depressed below the level of the epidermis itself. 

The aforementioned lips may be termed polar 
lips if they arise from the polar subsidiary cells 
or lateral lips if they arise from the lateral sub- 
sidiary cells. The polar lips are elongated par- 
allel to the long axis of the leaf and lie in a plane 
above the lateral lips which are elongated at 
right angles to the polar lips. This configuration 
of the lips causes great variation in the shape of 
the suprastomatal cavity in different planes of 
surface view, as noted by McClendon (1908). In 
our study five levels of change were seen for 
many species (Fig. 3): (A) the surface pore; (B) 
the level of the polar lips; (C) the level of the 
lateral lips; (D) the level of the chamber below 
the lateral lips or the inner aperture; (E) the level 
of the stomatal aperture. In some species fewer 
than five levels were observed, as in A. shawii 
ssp. goldmaniana where the polar lips are ab- 
sent. 


373 


FiGurRE 3. Levels of change in surface view of the Agave 
stomatal complex. A, surface pore; B, view of the polar lips; 
C, the lateral lips; D, the inner aperture; E, the stoma. 


The epidermal cells which surround the sto- 
matal complex may also possess lips or papillae, 
but these are less definite in form and number 
than those associated with the subsidiary cells; 
7-11 such cells surround the stomatal complex. 
Variations in the nature and development of the 
epidermal papillae or cuticle account for the vari- 
ation in the texture of leaf surfaces in Agave. 
The higher the papillae, the more asperous the 
leaf surface. Epidermal cells in general are either 
isodiametric (most species) or elongate (A. sob- 
ria ssp. sobria, A. sobria ssp. roseana, A. sobria 
ssp. frailensis, A. vizcainoensis , and A. subsim- 
plex), with end walls either transverse or 
oblique. 

The margin of the surface pore is of particular 
interest as it is formed in two different ways. In 


374 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 16 


avellanidens 


gigantensis 


subsimplex 


goldmaniana 


FiGureE 4. Variation in margins of the surface pore in eight 
species of Agave. 


most Deserticolae and all Campaniflorae, the 
surface pore is formed by the confluent cuticles 
of the cells surrounding the stomatal complex 
(in Fig. 4 see A. cerulata, A. moranii, A. avel- 
lanidens, and A. aurea). In some species the 
margin may appear irregular due to the presence 
of epidermal papillae (A. deserti) or, as in A. 
moranii where papillae are absent, the margin 
may appear entire. In the Umbelliflorae and in 
some Deserticolae, the surface pore is formed 
by the polar and lateral lips themselves, and the 
margin thus formed is entire (in Fig. 4 see A. 
subsimplex and A. shawii ssp. goldmaniana). 

In several species (Table 1) a peculiar and dis- 
tinctive ovoid structure can be seen at the level 
of the inner aperture. This appears to be due to 
a greater thickening of the cell wall of the polar 
and lateral-subsidiary cells adjacent to the inner 
aperture. We refer to this collarlike structure as 
a. Tim: + 


Stomata are so numerous in A. deserti and A. 
cerulata that the suprastomatal chambers tend 
to run together laterally forming a compound 
suprastomatal chamber or channel. A different 
situation is found in A. sobria where the margins 
of the surface pores tend to run together, aligned 
with the leaf axis rather than transversely. When 
‘‘channelling’’ is present, it is here referred to 
as transverse or axial channelling, respectively. 

For the most part stomata are evenly distrib- 
uted on both surfaces of the leaf, but they tend 
to become more sparse near the very base of the 
leaf. Consistent with the findings of other work- 
ers (Weisner and Baar 1914), there are more sto- 
mata on the upper surface of the leaf than on 
the lower (Table 2). 

The following characteristics of the stomatal 
complexes are described for each species: 


1. Number of levels of change in different 
planes of surface view 

2. Shape and nature of surface pore 

3. Length and width of the stomatal complex 

4. Width of the polar and lateral lips 

5. Position of the polar and lateral lips 

6. Length and width of the inner aperture 

7. Length and width of the stoma 

8. Presence of channelling 

9. Presence of rim 


The following descriptions are given in the 
form of brief diagnoses. The species are ar- 
ranged alphabetically under the sections. Sum- 
maries of epidermal characters are given in Ta- 
bles | and 2. Figures 5 to 50 show surface views 
of the Agave stomatal complex (photos all 
+ x200), stomatal ideographs, and rainfall sil- 
houettes of species habitats. The ideographs 
represent the relative sizes and shapes of the 
four subsidiary cells (white rectangles) and the 
sizes and positions of the polar lips (black rec- 
tangles). Sources of materials used are cited by 
collection numbers at the end of each paragraph. 


DESCRIPTION OF STOMATAL COMPLEXES 
Deserticolae 


Agave avellanidens Trelease (Figs. 5 & 14) 
Five planes in surface view, margin scalloped, 
extending out over the lateral subsidiary cells. 
Stomatal complex 105—130u long, 75—90u wide, 
oval to roundish in outline; polar lips 2, broad, 
extending towards one another but not touching, 
42-50 wide; lateral lips 2, apart or touching, 


a 


GENTRY & SAUCK: STOMATAL COMPLEX IN AGAVE 375 
TABLE 1. COMPARISONS OF EPIDERMAL CHARACTERS OF Agave. 
ee Se eS a ee 
Cuticle’ Stomatal? Position No. planes 
of polar pid. cell in surface 
Roughness Thickness Marg. Rim Chan. lips* elong.* view 
DESERTICOLAE 
avellanidens + ++ IR - + A + 5 
c. cerulata +4++ +++ IR = + A-T 4 5 
c. nelsonii +4++ +++ IR = =f A-T + 5 
c. subcerulata +++ +++ IR = + A + 5 
d. deserti +++ +++ IR = =f i) ar 5 
d. pringlei +++ +++ IR - + A-T + 5 
d. simplex +++ +++ IR - + T-O + 5 
gigantensis ar + ID = + A-T ar 5 
margaritae ++ +++ IR - + A + 5 
mckelveyana + ++ IR = * O +F 5 
moraniti + ++ ID + + A ++ 4-5 
s. frailensis + + IR + + A ++ 4-5 
5. roseana + + IR + + A ++ 5 
s. sobria +++ +4++ IR - + A ++ 5 
subsimplex ++ ++ IR + - A +44 4 
vizcainoensis + + E + - A ++ 3) 
CAMPANIFLORAE 
aurea + ++ IR = ‘4 A te 5 
capensis ++ aor IR = a A-T + 5 
promontorit “ft +++ IR = “e O + 5 
UMBELLIFLORAE 
s. goldmaniana + ft E 4 = = + 3 
sebastiana + spar E + = = ++ 3 
s. shawii + ++ E + - - + 3 
1 + = smooth or thin, +++ = rough or thick. 
2 1R = irregular, ID = indefinite, E = entire, — = absent, + = present, * = very occasional. 


3 A = polar lips apart, T = lips touching, O = lips overlapping. 


4 +4 = relative elongation. 


35-45 wide; inner aperture blunt oval, 46-60 
long, 28-35 wide; stoma 3040p long, 15—20u 
wide; channelling absent or occasional, trans- 
verse; surface smooth with small ridges of cutin, 
appearing slightly striate. Rim absent. (Gentry 
11929) 


Agave cerulata ssp. cerulata Trelease (Figs. 6 
& 15) 

Five planes in surface view, the margin of the 
surface pore irregular and extending out beyond 
the lateral cells. Stomatal complex 115-155 
long, 85-100 wide; polar lips 2, 50-60 wide, 
separated to touching, broadest at base, lateral 
lips 2, 40-63 wide, separated to overlapping; 
inner aperture rectangular 30-35 long and 31- 
40 wide. Stoma 33-40p long, 18-20u wide. 
Surface papillate and rough to touch; channel- 
ling present, transverse. Rim absent. (Gentry 
Garden, Murrieta, California) 


Agave cerulata ssp. nelsoni (Trelease) Gentry 
(Figs. 7 & 16) 

Similar to the above taxon, five planes in sur- 
face view, margin irregular and extending out 
beyond the lateral subsidiary cells. Stomatal 
complex 128-142 long, 90-100u wide, slightly 
longer than wide; polar lips 2, prominent, broad- 
est at base, apart or touching, 55-75 wide; lat- 
eral lips 2, touching to overlapping, 45-55 
wide: inner aperture elongate, 70-80 long, 23— 
29 wide. Stoma 34-38 long, 19-224 wide. 
Channelling occasional, transverse, surface pa- 
pillate. Rim absent. (Gentry 10373) 


Agave cerulata ssp. subcerulata Gentry (Figs. 
8 & 17) 

Five planes in surface view, margin irregular 
and extending out beyond the lateral subsidiary 
cells. Stomatal complex 140-180 long, 90—100u 
wide, more or less roundish in outline; polar lips 


376 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 16 
TABLE 2. STOMATAL MEASUREMENTS (MICRONS) OF Agave. 
Av. L&W Width Width Av. L&W Av. L&W Av. L&W Stomata/mm? 
stomate polar lat. inner lateral polar Av. L&W 
Taxa complex lips lips aperture cells cells stoma U L 
DESERTICOLAE 
avellanidens 120 x 80 45 39 —_— 66 x 21 37 x 42 _ 48 
c. cerulata 140 x 95 55 48 33.035 66 x 27 51 x 45 36 x 19 52 33 
c. nelsonii 132 x 96 67 53 (5x26 58 x 28 35 x 50 36 x 19 37 31 
c. subcerulata 150 x 88 45 42 59 x 27 565622 40 x 51 pH >< Ali 45 
d. deserti 128 x 82 46 44 — 49 x 18 30 x 38 — 52 34 
d. pringlei 110 x 105 45 38 47 x 18 48 x 22 32 x 42 28 x 10 61 
d. simplex Bil se ay 47 31 35 x 19 65 x 24 B25) — 
gigantensis 142 x 84 39 36 41 x 21 59 x 20 38 x 41 — 52 
margaritae 135 x 87 42 37 54 x 20 63 x 35 40 x 39 —_ 36 
mcekelveyana 114 x 80 35 23 37 x 20 47 x 22 33 x 51 — 23 
moranii 134 x 76 28 42 58 x 27 67 x 17 32 x 40 36 x 17 41 
Ss. frailensis 197 x 107 40 53 T2EXe38 70 x 36 71 x 46 35 x 14 30 
5. roseana 197 x 78 45 49 63 x 25 85 x 28 73 x 36 33 x 14 34 30 
s. sobria 153 x 91 48 37 — 71 x 35 48 x 49 = 36 
subsimplex 171 x 64 32 44 68 x 21 59 x 15 66 x 35 31x 8 42 
vizcainoensis 163 x 70 29 56 68 x 26 66 x 15 54 x 33 31 x 11 38 
CAMPANIFLORAE 
aurea 152 x 85 47 36 54 x 26 64 x 22 42 x 44 38 x 18 18 12 
capensis 125 x 80 33 38 47 x 23 41 x 21 38 x 34 34 x 13 31 19 
promontorii 126 x 84 67 40 43 x 29 65 x 25 38 x 47 — 21 18 
UMBELLIFLORAE 
s. goldmaniana 150 x 84 — 67 — 69 x 15 40 x 43 41 x 18 42 
sebastiana 135 x 76 _ 52 59 x 28 64 x 20 48 x 39 36 x 21 40 
s. shawii 142 x 78 = 53 43 x 41 72 x 20 48 x 43 39 x 22 47 36 


2, prominent, broadest at base, apart, 40-S50u 
wide; lateral lips 2, touching, 30—SOu wide; inner 
aperture somewhat irregular, generally oblong, 
54-65 long, 19-30u wide. Stoma 25-30p long, 
10-I5m wide. Channelling present, transverse; 
surface papillate. Rim absent. (Based on Gentry 
23175, 11892) 


Agave deserti Engelman ssp. deserti (Figs. 9 & 
18) 

Five planes in surface view, margin irregular 
and extending out over the lateral subsidiary 
cells. Stomatal complex 110-130 long, 80-90u 
wide, somewhat smaller than the cerulata type 
but similar to it. Polar lips 2, prominent, broad- 
est at base, touching or overlapping, 35—SOu 
wide; lateral lips 2, 45-50u broad, touching or 
overlapping; inner aperture 31-35 long, 17—20u 
wide rectangular. Channelling present, trans- 
verse; surface rough the papillae somewhat 
more acute than cerulata. Rim absent. (Gentry 
19741) 


Agave deserti ssp. pringlei (Engelman ex Tre- 
lease) Gentry (Figs. 10 & 19) 

Five planes in surface view, margin irregular 
as in subspecies deserti; stomatal complex 100- 
135p long, 85—115m wide, whole complex round- 
ish; polar lips 2, prominent, broadest at base, 
touching, apart, or overlapping, 38—SOu wide; 
lateral lips 28-40u wide, touching; inner aper- 
ture 40-54 long, 1S—22@ wide, somewhat con- 
stricted toward the center, stoma 22-30 long, 
10-11 wide; channelling present, transverse (as 
many as 6 stomata in a chain were observed); 
surface rough. Rim absent. (Gentry 19959, 
16723) 


Agave deserti ssp. simplex (Trelease) Gentry 
(Figs. 11 & 20) 

Five planes in surface view, margin highly ir- 
regular, much as in A. deserti; stomatal complex 
13lu long, 1124 wide, polar lips 47m wide, 
closed, lateral lips 2, 31 wide touching, inner 
aperture and stoma obscure, the cuticle very 


GENTRY & SAUCK: STOMATAL COMPLEX IN AGAVE 377 


FIG. 5 FIG. 8 FIG. 11 - 


A. avellanidens A. c. subcerulata A. d. simplex 

J D aj D Z D 
El Arco, B.C. San Ignacio, B.C. Aquila, Ariz. 

FIG. 6 FIG. 9 FIG. 12 I-I 

A. cerulata A. deserti deserti A. gigantensis 

J D J D J D 
Punta Prieta, B.C. Borrego Park, Calif. San Javier, B.C. 


FIG. 7 


FIG. 10 FIG. 13 
A. c. nelsonil A. d. pringlei A. margaritae 
J D J D J D 
sa Pe oe. Seeman —__ 
San Fernando, B.C. Valle Trinidad, B.C. Bahia Magdalena, B.C. 


FiGuRES 5 To 13. Ideographs of the Agave stomatal complexes with rainfall silhouettes of Agave habitats. Fig. 5. A. 
avellanidens. Fig. 6. A. cerulata cerulata. Fig. 7. A. cerulata nelsonii. Fig. 8. A. cerulata subcerulata. Fig. 9. A. deserti deserti. 
Fig. 10. A. deserti pringlei. Fig. 11. A. deserti simplex. Fig. 12. A. gigantensis. Fig. 13. A. margaritae. 


thick, surface papillate, channelling present, 160m long, 80-90 wide; polar lips 2, bulbous, 
transverse. Rim absent. (Gentry 23404) 32-46 wide, mostly slightly apart or touching, 
Agave gigantensis Gentry (Figs. 12 & 21) level with the leaf surface and continuous with 

Four levels in surface view, the margin of the the surface cutin, aperture formed at this level 
surface pore obscure; stomatal complex 116— perpendicular to the long axis of the leaf, lateral 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41. No. 16 


FIGURES 14 To 22. 


Surface views of the Agave stomatal complexes. Fig. 14. A. avellanidens. Fig. 15. A. cerulata cerulata. 
Fig. 16. A. cerulata nelsonii. Fig. 17. A. cerulata subcerulata. Fig. 18. A. deserti deserti. Fig. 19. A. deserti pringlei. Fig. 20. 
A. deserti simplex. Fig. 21. A. gigantensis. Fig. 22. A. margaritae. 


GENTRY & SAUCK: STOMATAL COMPLEX IN AGAVE 


lips below, 32—40u wide, overlapping; inner ap- 
erture 40-50u long, 20-264 wide; stoma ob- 
scure; channelling occasional, transverse, sur- 
face smooth. Rim absent. (Gentry 10237, 23320, 
Barclay & Argueles 1990) 


Agave margaritae Brandegee (Figs. 13 & 22) 
Five planes in surface view; margin of surface 
pore undulate and extending out beyond the lat- 
eral subsidiary cells; stomatal complex 115- 
160u long, 82-1004 wide, somewhat elongate; 
polar lips 2, 39-45u wide, slightly constricted 
toward base, apart; lateral lips 2, slightly over- 
lapping, 35—40u wide; inner aperture 50—60u 
long, 18-224 wide; stoma obscure; channelling 
occasional, transverse, general surface some- 
what rough. Rim absent. (Gentry et al. 11903) 


Agave mckelveyana Gentry (Figs. 23 & 32) 

Five planes in surface view; the margin of the 
surface pore irregular with 4-6 marginal pa- 
pillae, extending over the lateral subsidiary 
cells; stomatal complex 105-125 long, 75-85 
wide; polar lips 2, overlapping, 32-39 broad, 
lateral lips 2, overlapping, 21-23 broad, polar 
lips distinctly broader than the lateral lips; inner 
aperture 32-40u long, 19-22 wide; channelling 
very occasional, transverse; surface papillate. 
Rim absent. (Gentry & Ogden 9961) 


Agave moranii Gentry (Figs. 24 & 33) 

Four to five levels of change in surface view, 
the margin of the surface pore obscure or weakly 
developed, elongate; stomatal complex 122- 
150u long, 72-80 wide; polar lips 2, narrower 
than the lateral lips, far apart, 24-30 wide; lat- 
eral lips 2, touching or overlapping, more con- 
spicuous than the polar lips, 40-43 wide; inner 
aperture 55—60u long, 25—-30u wide, oblong to 
oval; stoma 32—-40u long, 15—18u wide; chan- 
nelling occasional, transverse; surface relatively 
smooth, cutin in small ridges. Rim present. 
(Gentry 23287) 


Agave sobria ssp. frailensis Gentry (Figs. 25 & 
34) 

Four to five planes in surface view; margin of 
the surface pore somewhat obscure, elongate, 
tending to connect with margins of adjacent sto- 
mates; stomatal complex elongate, nearly two 
times as long as broad, 185-203 long, 98-115 
wide; polar lips 2, not well-developed, far apart, 
30-45 broad; lateral lips broad, prominent, 
touching or overlapping, 50-64 wide, configu- 
ration of polar and lateral lips resembling that of 


379 


A. moranii; inner aperture oblong oval, 65—70u 
long, 15—20u wide; stoma 25-30 long, 13-16 
wide; surface essentially smooth, slightly striate; 
channelling present, axial. Rim present. (Gentry 
11257, 11264) 


Agave sobria ssp. roseana (Trelease) Gentry 
(Figs. 26 & 35) 

Five planes in surface view; margin elongate, 
undulate, tending to connect with margins of 
adjacent stomates; stomatal complex 140-175u 
long, 70-79 wide; polar lips 2, far apart, 42- 
47 wide; lateral lips 2, touching or overlapping 
slightly; 30-45 wide; inner aperture 62-73 
long, 20-22 wide; stoma 30-31 long, 18-20u 
wide; channelling present, axial; surface smooth, 
papillae visible but short. Rim present. (Gentry 
11277) 


Agave sobria ssp. sobria Brandegee (Figs. 27 & 
36) 

Five planes in surface view; margin of the sur- 
face pore irregular, elongate, and with some ten- 
dency to pass over the lateral subsidiary cells, 
this frequently not as marked as in A. cerulata 
and A. deserti; stomatal complex 137-175 long, 
90-107 wide; polar lips 2, prominent, generally 
apart, sometimes touching, 40—S5y wide; lateral 
lips 2, touching or overlapping slightly, 30-44 
wide; inner aperture and stoma obscure; surface 
lumpy, channelling axial and transverse. Rim 
absent. (Gentry 12387, 11811) 


Agave subsimplex Trelease (Figs. 28 & 37) 

Four levels in surface view; the polar and lat- 
eral lips forming a keyhole-shaped surface pore 
with an entire margin; stomatal complex dis- 
tinctly elongate, 150-190u long, 58-72 wide; 
polar lips 2, obscure, far apart, 28-34 wide; 
lateral lips 2, prominent, 38—SOu wide, apart; 
inner aperture 60-80 long, 15—25m wide, elon- 
gate, sometimes attenuated at the ends; stoma 
26-34u long, 7-9 wide; surface generally 
smooth, but with small ridges of cuticle running 
parallel with the long axis of the leaf; rim pres- 
ent. (Gentry 10217) 


Agave vizcainoensis Gentry (Figs. 29 & 38) 
Five planes in surface view, the margin of the 
surface pore elongate, nearly entire, slightly lon- 
ger and broader than the stomatal complex; sto- 
matal complex 160-172 long, 65-73 wide; po- 
lar lips 2, narrower than the lateral lips, 26-31 
wide, constricted toward base, far apart, never 
touching; lateral lips 2, broad, 54-60u wide, 


380 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 16 


FIG. 26 
FIG. 29 
FIG. 23 
A. mckelveyana A. s. roseana A. vizcainoensis 
J D J D J D 
oe ee | = —_—_ i+; 
Kingman, Ariz. La Paz, B.C. Punta Abreojos, B.C. 
FIG. 24 i oe FliGengo 
A. moranii A. sobria A. fortiflora 
J D J D J D 
Valle Trinidad, B.C. Comondu, B.C. Pitiquito, Sonora 
FIG. 28 
FIG. 25 eee 
A. s. frailensis A. subsimplex A. aurea 
J 2 D J D Ay Z D 
San Jose del Cabo, B.C. Carrizal, Sonora San Javier, B.C. 


FIGURES 23 TO 31. Ideographs of the Agave stomatal complexes with rainfall silhouettes of Agave habitats. Fig. 23. A. 
mckelveyana. Fig. 24. A. moranii. Fig. 25. A. sobria frailensis. Fig. 26. A. sobria roseana. Fig. 27. A. sobria sobria. Fig. 28. 
A. subsimplex. Fig. 29. A. vizcainoensis. Fig. 30. A. fortiflora. Fig. 31. A. aurea. 


GENTRY & SAUCK: STOMATAL COMPLEX IN AGAVE 


FicurEs 32 To 40. Surface views of the Agave stomatal complexes. Fig. 32. A. mckelveyana. Fig. 33. A. moranii. Fig. 34. 
A. sobria frailensis. Fig. 35. A. sobria roseana. Fig. 36. A sobria sobria. Fig. 37. A. subsimplex. Fig. 38. A. vizcainoensis. Fig. 
39. A. fortiflora. Fig. 40. A. aurea. 


382 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 16 


FIG. 43 FIG. 45 
FIG. 41 
A. capensis A. s. goldmaniana A. shawii 
J D J D J D 
Me ee ————_ | 
Cabo San Lucas, B.C. Punta Prieta, B.C. Ensenada, B.C. 
FIG. 42 FIG. 44 
A. promontorii 
A. sebastiana 
J 
J D 
Ei Mi 


Sierra Laguna B.C. 


FIGURES 41 To 45. 


Isla Cedros, B.C. 


Ideographs of the Agave stomatal complexes with rainfall silhouettes of Agave habitats. Fig. 41. A. 


capensis. Fig. 42. A promontorii. Fig. 43. A. shawii goldmaniana. Fig. 44. A. sebastiana. Fig. 45. A. shawii shawii. 


close together but not touching; inner aperture 
65-72 long, 22—30u wide, stoma 35-39 long, 
10-15 wide; channelling absent, surface 
smooth. Rim present, distinct. (Gentry 10339, 
7469) 


“Agave fortiflora Gentry (Figs. 30 & 39) 

Five layers in surface view; margin irregular 
due to cuticular papillae, similar to many De- 
serticolae in general aspect. Stomatal complex 
105-134 long, 83-99 wide; polar lips promi- 
nent, larger than the lateral lips, open or touch- 
ing, 35-45 wide, lateral lips 23-30 wide 
touching or overlapping; inner aperture 43-45 
long, 14-20u wide; stoma 25-30 long, 10-15u 
wide; channelling occasional, some polar con- 
nections; surface smooth but papillae visible. 
Rim absent. (Gentry 11630, 19808) 


Campaniflorae 


Agave aurea Brandegee (Figs. 31 & 40) 
Five planes in surface view; margin somewhat 


* Not aligned as to group. 


irregular and extending out beyond the lateral 
subsidiary cells, not continuous with the polar 
lips; stomatal complex 145-160u long, 80-92u 
wide, oval in outline; polar lips 2, conspicuous, 
constricted at base, apart, 40-50 wide; lateral 
lips 2, touching, 35-37 wide; inner aperture 47— 
60u long, 20-30 wide, oval; stoma 34—40u 
long, 15-18 wide; channelling occasional, 
transverse; surface smooth to slightly wrinkled, 
papillae when visible, small; cuticle distinctly 
yellow after staining. Rim absent. (Gentry 
12338) 


Agave capensis Gentry (Figs. 41 & 46) 

Five planes in surface view; margin as in A. 
aurea, not continuous with the polar lips; sto- 
matal complex 110—-135m long, 75-85u wide, 
oval; polar lips 2, apart, overlap occasionally, 
30-35 wide; lateral lips 2, mostly overlapping, 
larger than the polar lips, 35—40u wide; inner 
aperture 42—S50u long, 20-28 wide; stoma 32-— 
36m long, 11-15 wide; channelling occasional, 
transverse; surface smooth with some cutinous 
ridges; rim absent. (Santa Barbara Botanical 
Garden) 


GENTRY & SAUCK: STOMATAL COMPLEX IN AGAVE 


383 


Ficures 46 To 50. Surface views of the Agave stomatal complexes. Fig. 46. A. capensis. Fig. 47. A. promontorii. Fig. 48. 
A. shawii goldmaniana. Fig. 49. A. sebastiana. Fig. 50. A. shawii shawii. 


Agave promontorii Trelease (Figs. 42 & 47) 

Five planes in surface view; margin very ir- 
regular with as many as 8 cuticular papillae ob- 
scuring the lips below; stomatal complex 110- 
145u long, 80-874 wide; polar lips 2, broad, 
overlapping, 60-72 wide (nearly 2x those of 
capensis); lateral lips 2, touching, 35—45p wide, 
narrower than the polar lips; inner aperture 40— 
54u long, 28-33u wide, square to rectangular, 
stoma obscure, channelling occasional, trans- 
verse, surface more or less smooth; rim absent. 
(Gentry 11218) 


Umbelliflorae 


Agave shawii ssp. goldmaniana (Trelease) Gen- 
try (Figs. 43 & 48) 


Three levels of change in surface view, the 
surface pore formed by the lateral lips, entire, 
keyhole-shaped; stomatal complex elongate, 
140-190 long. 68-88 wide, polar lips absent, 
lateral lips prominent, apart or touching, 46-75 
wide; inner aperture 65-82 long, 37-42u wide, 
stoma 40-45 long, 16—20u wide; channelling 
absent, surface smooth, papillae absent. Rim 
present. (Desert Botanical Garden) 


Agave sebastiana Greene (Figs. 44 & 49) 

Not distinguishable from ssp. goldmaniana. 
Three levels of change in surface view; stomatal 
complex 110-160 long, 72-82 wide, polar lips 
essentially absent, lateral lips prominent, 40- 
55m wide, open; inner aperture oval, 56-634 
long, 27-304 wide; stoma 35-36 long, 18-25 


20 
i 
22 = 1\ 
bal 1 
ses ea 
20 = } 
1 \ 
=> (ee 
< i 
18 < | \ 
m2 = ! : 
° ! \ 
= 164105 | ct 
=< 1 \ / Se e 
' eee . 
a 14 x ; 
= I 
=< 1 
=_ xy 
12 
=x 
_ 5 ® 
< 
= 10 
o 
— 
| | | 
al <q 4 < <q 2) = 2) q 
<2 ee.) ES Gi ee ee 
= 28) &) .& | w & 
f& ae oo) wia «a 
Peja > oo = 6 
zaneqaqge?®ewY®2zwyw 
Od w Ss w z a < 
Se SE a (Og = 
qt w 3 = 
Ss o wi 
o 2 AN = 
o nD = 
> < 
FIGURE 51. 
A. margaritae 135 X 87 = 11,745u”) and the stomatal index (I = 


wide; no channelling, surface smooth. Rim pres- 
ent. (Moran 15142, 17430; Beauchamp 2095) 


Agave shawii Engelman ssp. shawii (Figs. 45 & 
50) 

Very similar to the above subspecies. Sto- 
matal complex elongate, 137—150u long, 67-83 
wide; polar lips absent or rudimentary, lateral 
lips prominent, 45-60 broad; inner aperture 
42-60u long, 30-35 wide, stoma 34-39 long, 
20-214 wide; channelling absent, surface 
smooth. Rim present. (Gentry 10397) 


DISCUSSION AND CONCLUSIONS 


Certain conclusions can be drawn regarding 
the value of the stomatal cell complex for sys- 
tematic studies. We found, as did McClendon 
(1908), that in many taxa the surface aperture 
(level of the polar lips) of the suprastomatal pas- 
sage is usually elongated at right angles to the 
long axis of the leaf, and farther down (level of 


MORANII [4] 
SUBSIMPLEX [___ 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 16 


SO 


2 
° 
MM PRECIPITATION 


AUREA 


= 
< 
x 
7) 


SIMPLEX ‘(2-22 2] 
FRAILENSIS [sd 


CAPENSIS 
GIGANTENSIS 
MCKELVEYANA 
PROMONTORII 


Average annual rainfall of Agave habitats compared with stomatal surface areas (data from Table 2; for example, 


S/(E + S) x 100), after Wicks (1935). 


the lateral lips) it is elongated parallel to the long 
axis of the leaf. Blunden et al. (1973) observed 
the reverse in A. ellemeetiana. We have no ma- 
terial of this species, but we have observed in 
others that the surface pore may appear elon- 
gated parallel to the long axis of the leaf if the 
polar lips are absent or rudimentary—as in all 
the taxa of the Umbelliflorae and in A. subsim- 
plex. 

The presence of four cells surrounding the 
guard cells appears to be a constant character 
in over 80 species of Agave, if we include our 
data with those of Blunden et al. (1973). That 
the stomatal complex is only morphologically 
tetracytic must be emphasized, as our observa- 
tions refer to the appearance of the stomata 
when fully differentiated. 

A puzzling feature described by Blunden et 
al. (1973) for several agavaceous genera and by 
Blunden and Binns (1970) for Yucca glauca is 


GENTRY & SAUCK: STOMATAL COMPLEX IN AGAVE 


the presence of beaded cell walls. We have ob- 
served no such structure in our work with 
Agave. It is possible that the epidermal portion 
containing the beaded cell walls is not obtained 
in leaf peelings. 

Taxonomic use of the epidermal characters 
can be considered on the sectional, specific, and 
subspecific levels. The stomata of the Umbelli- 
florae, Campaniflorae, and Deserticolae are all 
distinct from one another, as represented by 
their respective ideographs and by their surface 
views (Figs. 5—S0). 

Most of the stomatal elements or characters 
are common to all three groups, but a few are 
lacking in one or another group. While stomatal 
rims are lacking in the Campaniflorae, they are 
present in the Umbelliflorae, and absent or pres- 
ent in the Deserticolae. While the lack of a single 
element may not be sufficient to support group 
status, it becomes notably significant when sup- 
ported by other characters. Variations in the 
lips, margins, and number of changes in levels 
of surface view of the stomata are especially 
useful in distinguishing groups. The Umbelli- 
florae and Campaniflorae are small distinct 
groups, but the Deserticolae is large, wide- 
spread, and complex in the variability displayed 
by its seventeen taxa. 

In the species of the Deserticolae—a fairly 
coherent group macromorphologically—we have 
found a surprising amount of diversity in sto- 
matal structure. Major features of the stomatal 
complex were found to be constant for a spe- 
cies, while variation between species was often 
apparent. Species such as A. cerulata and A. 
deserti possess thick cuticles with elaborate sur- 
faces, deeply depressed stomata, and conspic- 
uous polar and lateral lips which tend to obscure 
the suprastomatal chamber. In contrast, A. sob- 
ria frailensis, A. sobria roseana, A. subsimplex, 
and A. vizcainoensis possess relatively smooth 
surfaces, with stomata not deeply depressed, 
and poorly developed polar lips leaving the su- 
prastomatal chamber below ‘“‘unprotected.”’ 

In order to distinguish species of the Deser- 
ticolae, it was found necessary in most cases to 
use a combination of leaf-surface characters 
with the variations in the lips, margins, and 
number of changes in the levels in surface view 
of the stomata. For instance, A. cerulata and A. 
deserti can be distinguished from A. sobria ro- 
seana and A. sobria frailensis by the presence 


385 


of a rim in the latter two subspecies. A. subsim- 
plex can be distinguished from all other Deser- 
ticolae in having four planes in surface view, a 
rim, and the margin of the surface pore formed 
by the lateral lips. Other species were not found 
separable by stomatal characters, as the two 
complexes represented by A. deserti and A. cer- 
ulata, both of which have highly variable over- 
lapping subspecific forms. 

In the Campaniflorae, A. aurea can be distin- 
guished from A. capensis in that the polar lips 
of aurea are 40—S0mu wide, while those of capen- 
sis are 30—35u wide. A. promontorii is distin- 
guished from the first two not only by its wider 
polar lips (60-72), but also by its thicker cuticle 
and strongly overlapping polar lips. 

In the Umbelliflorae the polar lips are absent 
and the margin of the surface pore is entire. The 
stomatal complex is very uniform and thus can- 
not be used for distinguishing the subspecies. 
The stomatal-cell complex, however, is suffi- 
ciently distinct to separate this group readily 
from the Deserticalae and the Campaniflorae. A 
species with uncertain affinities, A. fortiflora, 
was compared with the Umbelliflorae, as it bears 
some resemblance to this group in floral mor- 
phology. It had been placed there previously by 
Gentry (1972). However, the stomatal complex 
of A. fortiflora is found to bear no resemblance 
to that of the Umbelliflorae and it has been re- 
moved from that group. 

Microscopic examination of the epidermis is 
frequently of particular value in the identifica- 
tion of species. This is true with isolated leaf 
specimens, which in form or armature may be 
atypical of a given species. For instance, Howell 
10660, a leaf specimen from San Bartoleme Bay, 
resembles in size and armature A. margaritae, 
but examination of the stomata shows it to be 
conspecific with A. vizcainoensis. As another 
example; the latter taxon, on gross morpholog- 
ical appearance and smooth cuticle, was first 
judged to be a part of A. gigantensis, but when 
stomatal structure was examinaed, A. vizcain- 
oensis was set aside as a separate species. 

It would be possible to classify Agave on ep- 
idermal criteria alone (cf. Muller 1909), but the 
relationships would appear very different from 
those obtained on the criteria of leaf and inflo- 
rescence morphology. A sound taxonomic sys- 
tem must be built on a broad range of criteria. 
Epidermal criteria are a valuable supplement to 


386 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 16 


classical morphology and to the newer guides of 
growth and breeding habits. 

The thick cuticles with elaborate surfaces, 
deeply depressed stomata, and conspicuous po- 
lar and lateral lips, which tend to obscure or 
cover the suprastomatal chamber, appear to be 
xeromorphic adaptations for desert survival. 
This observation is discussed below. 


Cuticular Structure and the Environment 


The cuticle of Agave is a remarkable structure 
forming a translucent envelope over the func- 
tional cells within the leaf. It appears to diffuse 
sunlight, insolate against extreme temperatures, 
protect against winds and mechanical abrasion, 
reduce transpiration, and support the succulent, 
heavy leaf as a whole. The unusual thickness 
and other characteristics of the cuticle suggest 
that it has evolved in response to the aridity of 
the environments in which most of the species 
occur. McClendon (1908) paid particular atten- 
tion to this theme and observed that the more 
xerophytic species of Yucca, Nolina, Dasyli- 
rion, and Agave had specialized stomatal struc- 
tures. He states that the suprastomatal passages 
are guarded by four lips, which with the asso- 
ciated subsidiary cells, narrow or close the pas- 
sage in several species, e.g., Yucca elata, Y. 
rostrata, Agave victoriae-reginae, A. schottii. 
Nolina, as well as some Agave species, have 
stomata aligned in deep grooves. These struc- 
tures, he noted, served by ‘‘protecting the sto- 
mata from the dry air’? (McClendon 1908:316). 

In Figures 5 to 50 we have provided rainfall 
silhouettes along with stomatal ideographs and 
surface views of the stomata. Rainfall data were 
extracted from the National Climatic Center 
(1973) for Arizona, and Hastings (1964a and 
1964b) for Mexico. With these we hope to call 
attention graphically to possible correlation be- 
tween stomatal morphology and rainfall pat- 
terns, the most critical environmental factor for 
desert plants. Detailed conmparisons between 
environmental factors and epidermal structure 
is beyond the scope of this study, but a few ob- 
servations can be made. 

The rainfall profiles show that all stations, ex- 
cept for one or two, and their nearby Agave pop- 
ulations are strictly desertic. One certain excep- 
tion is the Sierra Laguna station in the Cape 
District, with 747 mm of annual rainfall; it is the 
montane habitat of Agave promontorii, a species 


associated with forests. For several Baja Cali- 
fornia stations the available records are short 
term, 5—7 years, and therefore of limited reli- 
ability. The one for A. cerulata ssp. nelsonii, 
San Fernando on Sierra San Miguel, is obvious- 
ly rejectable because of its low figure of only 66 
mm for average annual rainfall. Generally, we 
can see no correlations between rainfall patterns 
and variations in stomatal structure. This is the 
case for annual rainfall, length of dry season, 
and for both winter-rainfall season and summer- 
rainfall season. The measurements of stomatal 
structures are also noncorrelative with rainfall 
patterns, as for instance, the average surface 
area of the stomatal complex (120u x 80 = 
9,600u for A. avellanidens, and so on for the 
rest) (Fig. 28). 

On the basis of rainfall data we do not find, 
as McClendon (1908) observed, that closure of 
the suprastomatal passage is a xeromorphic ad- 
aptation. A. promontorii, for instance, has the 
surface opening completely closed by the polar 
lips (Fig. 47), but its habitat is by far the most 
watered of all our subject species. Taxa of the 
A. shawii group are all characterized by lipless 
open stomatal surface openings (Figs. 48—50) in 
a smooth cuticle of moderate thickness, yet both 
sebastiana and goldmaniana occupy the drier 
habitats having less than 100 mm of annual rain- 
fall. However, these three taxa occupy the 
heavy fog belt along the outer peninsular coast. 
Their distinctive, elongate, lipless stomata may 
well reflect structurally the sustaining dew of the 
fog-desert. This apparent correlation is also sup- 
ported by the relatively open lips and shallowly 
depressed stomata in smooth cuticular surfaces 
observed in A. sobria ssp. frailensis, A. sobria 
ssp. roseana, and A. vizcainoensis (group De- 
serticolae), all of which are maritime dwellers 
where fog or high humidity are relatively fre- 
quent. However, A. moranii, with similar sto- 
matal structure, is an inland species on the very 
arid eastern slopes of the Sierra San Pedro Mar- 
tir. There are no fog or humidity data available 
to verify these observations. 

At this point we cannot argue for or against 
the concept of epidermal structure having de- 
veloped in response to desert aridity. It is ap- 
parent that we need to know much more about 
both structure and function as well as climato- 
logical and other factors distinguishing habitats. 
For instance, the habit of day closure—night 


GENTRY & SAUCK: STOMATAL COMPLEX IN AGAVE 


opening of stomates may be more significant in 
desertic evolution than the structures discussed 
above. This habit reduces transpiration, con- 
serves moisture, and contributes towards sur- 
vival in arid climates. Many succulents, includ- 
ing agaves, have been found with this physiologic 
habit, known as CAM (for Crassulacean Acid 
Metabolism). There is a rapidly growing litera- 
ture on this subject (see Mooney et al. 1974; 
Szarek and Troughton 1976). 

The agave cuticle appears to function like an 
air-conditioned greenhouse protecting the tend- 
er, living leaf tissue from desertic extremes. The 
general characteristics of this structure are com- 
mon to all agaves and some other related genera. 
However, the stomates of the two narrow en- 
demic groups, Umbelliflorae and Campaniflor- 
ae, have distinctive features. Just how the struc- 
ture and function of their stomates are correlated 
with their narrow environments is not clear at 
this writing. Nor is it clear how or if these struc- 
tures have any survival value for the species. 
Some ideas need testing, others appear as self- 
evident truths, and still others are transitional to 
other ideas. We hope that the idea about pro- 
tective xeromorphic stomates will stimulate fur- 
ther research. 


ACKNOWLEDGMENTS 


We thank Dr. James E. Canright, Department 
of Botany and Microbiology, Arizona State Uni- 
versity, for graciously making laboratory equip- 
ment and space available for slide preparation, 
and Dr. Donald J. Pinkava for his editorial com- 
ments on our manuscript. This study was made 
possible through National Science Foundation 
grant no. BMS74-24553. 


LITERATURE CITED 


BLUNDEN, G., AND W. W. BINNS. 1970. The leaf anatomy 
of Yucca glauca Nutt. Bot. J. Linn. Soc. 63(2):133-141. 

, Yt Y1, AND K. JEWeErRS. 1973. The comparative leaf 
anatomy of Agave, Beschorneria, Doryanthes and Furcraea 
(Agavacea). Bot. J. Linn. Soc. 66(2):157—159. 

GENTRY, HowARD Scott. 1972. The agave family in Sonora. 


387 


U.S. Dep. Agric. Agric. Handbk. 399. Pp. 1-195, maps, 

illus. 

1978. The agaves of Baja California. Occas. Pap. 
Calif. Acad. Sci. 130. Frontis. + 6 color pls., 61 figs., 
11 tables. 

HASTINGS, J. R. (ed.) 1964a. Climatological data for Baja 
California. Technical reports on the meteorology and cli- 
matology of arid regions, no. 14. Univ. Ariz. Inst. Atmos. 
Physics, Tucson, Arizona. 

. 1964b. Climatological data for Sonora and northern 
Sinaloa. Technical reports on the meteorology and clima- 
tology of arid regions, no. 15. Univ. Ariz. Inst. Atmos. 
Physics, Tucson, Arizona. 

JOHANSEN, DONALD A. 1940. Plant microtechnique. Mc- 
Graw-Hill, New York. 

McCLENDON, J. F. 1908. On xerophytic adaptations of leaf 
structure in Yucca, Agave, and Nolina. Am. Nat. 42:308- 
316. 

Mooney, H., J. H. TROUGHTON, AND J. A. BERRY. 1974. 
Arid climates and photosynthetic systems. Carnegie Inst. 
Wash. Yearbook 73:793-805. 

MULLER, CARL. 1909. Beitrage zur vergleichenden Anatomie 
der Blatter der Gattung Agave und ihrer Verwertung fiir die 
Unterscheidung der Arten. Bot. Ztg. 67(1):93-141. 

Napp-ZINN, K. 1974. Anatomie des Blattes, II. Blattanato- 
mie der Angiospermen. /n Encyclopedia of plant anatomy. 
Gebrider Borntraeger, Berlin. 

NATIONAL CLIMATIC CENTER. 1973. Climatography of the 
United States no. 81, Arizona. U.S. Dep. Commerce. 

SHAH, G. L., AND B. V. GopAL. 1969. Structure and devel- 
opment of stomata on the vegetative and floral organs of 
some Amaryllidaceae. Ann. Bot. (Lond.), n.s., 34:737-749. 

SINCLAIR, C. B., AND D. B. DUNN. 1961. Surface printing 
of plant leaves for phylogenetic studies. Stain Technol. 
36:299-304. 

Stace, C. A. 1966. The use of epidermal characters in phy- 
logenetic considerations. New Phytol. 65(3):304—-318. 

STEBBINS, G. L., AND S. KHUSH. 1961. Variation in the or- 
ganization of the stomatal complex in the leaf epidermis of 
monocotyledons and its bearing on their phylogeny. Am. J. 
Bot. 48:51—S9. 

SZAREK, S. R., AND J. H. TROUGHTON. 1976. Carbon isotype 
ratios in crassulacean acid metabolism plants. Plant Physiol. 
58:367-370. 

TOMLINSON, P. B. 1974. Development of the stomatal com- 
plex as a taxonomic character in the monocotyledons. Tax- 
on 23:109-128. 

Wicks, L. M. 1935. The anatomy of amaryllidaceous leaves. 
I. Stomatal distribution in Haemanthus and Brunsvigia. 
Ann. Bot. (Lond.) 49:493—S05. 

WIESNER, J. V., AND H. Baar. 1914. Beitrage zur Kenntnis 
der Anatomie des Agave-Blattes. Sitzungsber. Akad. Wiss. 
Wien, Math.-Naturwiss. K]., Abt. I, Bd. 123:679-714. 


iC wees | ‘)(\ ee | 
vu J be ; al ri a 


i) 


- 


: 2 


os 


pie AO tren o 


J ® 2 ) — nay 
: UP Go _ran | ee. es epic Toa’, 
——.  @ @e-4 ; =f..ie 


2 ta” @o), a5 - ani Dibeg ied may Or wie ; 
Am " tina Che 0 ee 
| on ee 4. O ve onan f Sof! oh! 
j bay - 7» oa ae 


: @ ) 2 eh oud 
; ree a 
; : ri - - , oy 


, 4 Ti) ey 
us Ti. a) im 


a 40 Oh Bie 


= i b wi 
Pa ure We nO has = 
7 0 Gel ME ol os «| 
Taha ON alla 


2 eo : ? a 
fue we 7 \esl hing! at 
sbi Seen 


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PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 
FOURTH SERIES 


Vol. XLI, No. 17, pp. 389-399; 7 figs. 


December 28, 1978 


ANATOMICAL NOTES ON AN ADULT MALE OF THE DEEP-SEA 
OPHIDIID FISH PARABASSOGIGAS GRANDIS 
FROM OFF CALIFORNIA 


Carl L. Hubbs 


Scripps Institution of Oceanography, University of California, San Diego, 
La Jolla, California 92093 


and 
W. I. Follett 
California Academy of Sciences, San Francisco, California 94118 


ABSTRACT: 


Anatomical notes on a large adult male of the deep-sea ophidiid fish Parabassogigas grandis 


(Giinther, 1877), the first known specimen of this species from off California, include description of the 
lateral-line system, the gill rakers on all arches, and the caudal-fin complex. The median rocker bone, 
precaudal vertebrae, gas bladder, and associated musculature, presumably constituting the sound-producing 


mechanism of the adult male, are discussed. 


A large fish (1,353 mm in total length, 16.5 kg 
in weight), collected off the Farallon Islands on 
22 September 1952, represents the only known 
specimen referable to the deep-sea ophidiid Par- 
abassogigas grandis (Gunther, 1877) from Cal- 
ifornia (Fig. 1). 

The fish was collected by commercial fisher- 
men Warren E. Beadle and R. G. Hamilton 
(aboard their albacore boat, ‘‘New Daisy’’), 
who had observed it floating some 40 miles west- 
southwest of Farallon Lighthouse, a well-known 
landmark 37 km southwest of Point Bonita at 
the entrance to San Francisco Bay. The approx- 
imate position, computed from U.S. Coast & 
Geodetic Survey Chart 5002 (1940), was 
37°30'N, 123°39'W, where the depth of water is 
2,926 m. The weather was flat-calm; the surface 
temperature of the water was 15.5°C. The fish, 


[389] 


which was still bleeding, with ‘‘air bubbles’’ un- 
der the skin, had been somewhat mutilated 
about the belly and dorsal fin by two or three 
blue sharks (Prionace glauca) and albatrosses 
that were observed ‘working on the fish.”” It 
was dead, presumably as a result of the clogging 
of its intestine by cestodes. 

On 24 September 1952, the fish was presented 
to the Department of Ichthyology of the Cali- 
fornia Academy of Sciences (catalog no. 25724). 
The specimen was recorded by Follett (1970:481). 


METHODS 


Counts and measurements were taken as by 
Hubbs and Lagler (1958:19-26). Gill-raker 
counts include all rudiments, however small. We 
follow the lepidological terminology of Lagler 
(1947) and the osteological terminology of 


390 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 17 


FIGURE 1. 


Parabassogigas grandis (Gunther), CAS 25724, standard length 1,270 mm, total length 1,353 mm, weight 16.5 


kg, from off the Farallon Islands, California. Photograph by Moulin Studios, San Francisco. 


Weitzman (1967). Counts of the median fin rays, 
and all vertebral characters, are from radio- 
graphs. Counts and measurements of the two 
sides are separated by a diagonal (/). 


SYNONYMY 


Sirembo grandis GUNTHER, 1877:437-438 (original descrip- 
tion; south of Yeddo [Tokyo], Japan).—JoRDAN, TANAKA, 
AND SNYDER 1913:404 (type-species of Bassogigas; refer- 
ence). 

Neobythites grandis: GUNTHER 1887:100-102, pl. 21, fig. A 
(redescription; holotype from south of Yedo, near Yoko- 
hama, CHALLENGER sta. 237; 1,875 fathoms (3,429 m); 
{trawled 17 June 1875; 34°37'N, 140°32’E; mud; bottom 
temp. 35.3°F (1.8°C); surface 73.0°F (22.8°C)]; data in brack- 
ets from Thomson 1880:57).—GooDE AND BEAN 1896:329 
(reference; compared with Bassogigas gillii) —GARMAN 
1899:391 (listed). —BRAUER 1906:403 (reference to type-lo- 
cality)—DoLLo 1906:174, 176 (reference). 

Bassogigas grandis: GOODE AND BEAN 1896:529 (type-local- 
ity after Gunther 1887).—JORDAN AND FOWLER 1902:758- 
759 (synonymy; description copied).—TANAKA 1908:1, 17— 
18, pl. 2, figs. 1A-1B (description; 1 specimen; Sagami 
Sea, Japan).—JORDAN, TANAKA, AND SNYDER 1913:404 
(soko-bozu [bottom priest]; reference; distribution).—Nor- 
MAN 1939:86 (comparisons; relationships).—NYBELIN 
1948:80, 81 (comparisons; in key)—KAMOHARA 1954:2 (in 
key).—NyYBELIN 1954:67 (distribution).—MATSUBARA 
1955:799 (soko-bozu; distribution) —GreyY 1956:207 (syn- 
onymy; distribution). 

Parabassogigas grandis: NYBELIN 1957:296, 298, 345 (rela- 
tionships; Parabassogigas, new genus, type-species, by 
original designation, Sirembo grandis Giinther; type-local- 
ity after Giinther 1887); fig. 31 (dentition).—EAGLE 
1969: 1680-1685 (description; 6 specimens off Newport, Or- 
egon, 2,800 m; 1 specimen off Tillamook, Oregon, 2,130 m); 
fig. 1 (351-mm female); fig. 2 (vomerine and posterior ba- 
sibranchial tooth patches); figs. 3-7 (left gill arch); figs. 8— 
9 (angle gill raker)—FoLLETT 1970:481 (caudal-ray count 


of Calif. Acad. Sci. no. 25724, discussed in this paper, and 
of Zool. Inst. Univ. Tokyo no. 39255, from off Katsu-ura, 
Japan). 


DESCRIPTION 


Measurements.—The following measure- 
ments, additional to those noted below, are ex- 
pressed in millimeters: Standard length 1,270; 
total length 1,353. Body depth at dorsal origin 
300, at anus (posterior edge) 240. Caudal pedun- 
cle least depth 13. Body thickness at pectoral 
base 154. Snout tip to dorsal origin 352, to anal- 
fin origin 614, to pectoral base (upper end) 252/ 
255, to left pelvic base 184. Anus (posterior 
edge) to caudal base 698. Head length 237/235. 
Snout length 72/72. Orbit, horizontal diameter, 
22/22. Interorbital width 97. Suborbital least 
width 31.5/32. Anterior nostrils, least distance 
between, 44. Posterior nostrils, least distance 
between, 77. Anterior nostril to posterior nostril 
14.5/16. Posterior nostril (upper end) to orbit 14/ 
13. Maxillary length 131/127, depth (greatest 
fleshy) 28/31. Preopercular groove to end of 
opercular membrane 61/65. Opercular spine 
(lower free edge) 28/29. Pectoral length 126/121; 
pectoral base depth 67/67. Left pelvic outer-ray 
length 81; inner-ray (tip broken off) width 2.4. 
Premaxillary tooth-patch length 87/87, width 
5/5. Vomerine tooth-patch anteroposterior length 
12, width 24. Palatine tooth-patch length 58/54, 
width 11/11. Dentary tooth-patch length 90/90, 
width 6/6. Anterior basibranchial tooth-patch 
length 36, width 10. Posterior basibranchial 
tooth-patch length 14, width 8. First superior 


HUBBS & FOLLETT: OPHIDIID FISH PARABASSOGIGAS GRANDIS 391 


pharyngeal tooth-patch length 21/21, width 8/8. 
Inferior pharyngeal tooth-patch length 42/42, 
width 8/8. 

Head.—Head rather short, smooth, with no 
suggestion of crests; its anterodorsal profile 
evenly rounded except for a faint convexity at 
snout. Interorbital strongly convex; its least ver- 
tical height above anterodorsal margin of orbit 
44 mm. Eye relatively small and far forward, 
with no externally evident differentiation of iris 
and pupil. No free orbital rim. Snout barely pro- 
duced above the large mouth; lower jaw slightly 
included. No barbels. Premaxillaries only slight- 
ly protractile. Maxillary extending 47 mm be- 
hind eye. Anterior nostril a low, thick tube; pos- 
terior nostril an open, crescentic slit. Anterior 
nostril separated by a groove, behind and below, 
from a row of 5 compressed papillae. This 
groove shallowly connected with a deep sulcus 
extending to rostral edge, becoming shallow at 
the edge. Anterior edge of sulcus a scaleless 
ridge running downward and forward and meet- 
ing at an angle of approximately 50° the lateral 
lobe of a scaleless rim that extends around front 
of snout. A flaplike papilla concealed under ros- 
tral fold opposite sulcus; between this papilla 
and midline of snout a larger, similar papilla. 
Membrane of opercle above continuous with an 
extremely narrow flap extending downward on 
shoulder girdle to pectoral base; membrane 
readily apparent near its end, but so reduced in 
height at midregion as to be scarcely percepti- 
ble. Opercular spine simple and deeply embed- 
ded, its tip (apparent in a dermal aperture) about 
25 mm anterior to fleshy opercular margin. No 
pseudobranchiae. Behind left 4th gill arch, a slit 
49 mm long; behind right, a slit 43 mm long. Gill 
membranes free from isthmus. Branchiostegals 
8 (4 + 4) on each side—this number usual in 
fishes formerly placed in the Brotulidae, rare 
among spiny-rayed fishes and their derivatives 
(Hubbs 1919:70). 

Dentition.—Villiform teeth cover the premax- 
illaries, vomer, palatines, dentaries, anterior and 
posterior basibranchials, 2nd and 3rd epibran- 
chials, all 3 superior pharyngeals, and the infe- 
rior pharyngeals. The individual teeth are blunt- 
ly conical, straight or somewhat curved, and of 
nearly uniform size; the largest is 0.8 mm in 
height and 0.3 mm in basal diameter. There are 
approximately 305 teeth in the vomerine tooth 
patch (Fig. 2, top), which resembles a triangle 
with nearly straight base, slightly concave sides, 


Dentition of Parabassogigas grandis, CAS 


FIGURE 2. 
25724. Top, premaxillaries, vomer, and palatines. Bottom, 
dentaries, anterior and posterior basibranchials, and inferior 
pharyngeals. Photographs by W. I. Follett. 


and parabolic vertex. The inner margin of each 
palatine tooth patch (Fig. 2, top) is concave an- 
teriorly. The basibranchials (Fig. 2, bottom) 
bear two median unpaired tooth patches (a larg- 
er anterior and a smaller posterior patch), well 
separated; they lack the intervening pair that is 
present in the basibranchials of Bassogigas as 
figured by Nybelin (1957: figs. 25-30, 50). The 
inferior pharyngeal patches (Fig. 2, bottom) are 
lanceolate, with apex directed posterolaterally. 
For dimensions of tooth patches, see Measure- 
ments. 


392 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 17 


FIGURE 3. 


First gill arch (left) of Parabassogigas grandis, 
CAS 25724. Top, outer (anterior) aspect. Bottom, inner (pos- 
terior) aspect. Photographs by W. I. Follett. 


Gill Rakers.—First arch, outer (anterior) se- 
ries 5 + 21/5 + 20. Left (Fig. 3, top): On upper 
limb 5 flat, finely dentigerous plates (Sth dou- 
bled). On lower limb (including angle) 5 stubby, 
clavate rakers approximately 11 mm long, 6 mm 
wide, 4 mm thick (with exceedingly minute bris- 
tles on tips), a small plate between 3rd and 4th 
and 4th and Sth rakers; then 8 structures, vary- 
ing irregularly from plates of diverse size to rak- 
ers, 8th a stubby, clavate raker 9 mm long; then 
6 large, more or less fused, finely dentigerous 
plates, nearly covering arch and forming an even 
edge opposite bases of gill filaments. Right: On 
upper limb 5 large plates. On lower limb (in- 
cluding angle) 7 stubby, clavate rakers approx- 
imately 10 mm long, a small plate in 4th, Sth, 


and 6th intervals; then 2 larger plates alternating 
with 2 unequal rakers; then 6 large, more or less 
fused, finely dentigerous plates. 

First arch, inner (posterior) series 4 + 15 / 
4+ 16. Left (Fig. 3, bottom): On upper limb all 
structures are plates. On lower limb they vary 
irregularly from plates to rakers; 2nd and 4th to 
11th are blunt rakers. Right: On upper limb all 
structures except 4th (a tiny rudiment) are 
plates. Those on lower limb vary from plates to 
rakers; 3rd and 5th to 12th are blunt rakers. 

Second arch, outer series 5 + 18/5 + 18. On 
this and all subsequent arches all structures (ex- 
cept as noted) are flat plates, closely juxtaposed, 
nearly covering arch and forming an even edge 
opposite bases of gill filaments. Left: On lower 
limb 8 blunt rakers alternating with large plates; 
then 3 blunt rakers grading into plates. Right: 
On lower limb 6 blunt rakers alternating with 
large plates; then 7 blunt rakers grading into 
plates. 

Second arch) inner series 3°+ 17/92): 
Left: On lower limb first 6 structures are blunt 
rakers alternating with small plates; 3rd, Sth, 
and 7th to 12th plates are cordate. Right: On 
lower limb 3rd to 8th structures are blunt rakers 
alternating with small plates; 7th and 9th to 1Sth 
plates are cordate. 

Third arch, outer series 4 + 16/3 + 16. Left 
and right: On lower limb 3rd and Sth to 11th 
structures are blunt rakers or at least somewhat 
carinate plates. 

Third arch; inner series:2 +. 12/2 2a 
On lower limb 4th to 9th plates are cordate. 
Right: On lower limb 4th to 7th plates are cor- 
date. 

Fourth arch, outer series 1 + 10/2 + 11. Left 
and right: On lower limb 2nd to 7th plates are 
slightly carinate, with blunt tips. 

Fourth arch, inner series 0 + 8 / 0 + 7. Left 
and right: The plates are somewhat convex dis- 
tally. 

Fifth arch 0 + 7/1 + 8. Left: On lower limb 
7 flat contiguous plates increase in size poste- 
riorly. Right: On upper limb is a small raker. On 
lower limb plates increase in size posteriorly. 

Otolith.—The right sagitta (Fig. 4, top, lateral 
aspect; Fig. 4, bottom, mesial) is subpentagonal, 
its dorsal margin slightly concave, its ventral 
margin strongly convex. It is 12.5 mm long, 10.8 
mm high, and 4.2 mm thick. The sulcus com- 
prises the ostium (largely the anterior part of the 
sulcus) and the cauda (the posterior part of the 


HUBBS & FOLLETT: OPHIDIID FISH PARABASSOGIGAS GRANDIS 393 


sulcus and a narrow elongate area above the os- 
tium). The dorsal margin of the sulcus is nearly 
a straight line except where the sulcus opens 
upward at about 45° toward the anterodorsal 
margin of the sagitta. The ventral margin curves 
downward to slightly beyond the middle of the 
sagitta, thence sharply upward to the ventral 
margin of the cauda (at the stricture). The great- 
est vertical distance from the dorsal margin of 
the sagitta to the dorsal margin of the sulcus is 
2.8 mm; the distance from the ventral margin of 
the sulcus to the ventral margin of the sagitta, 
5.0 mm. The rounded area below the anterodor- 
sal opening of the sulcus (presulcal trough) con- 
stitutes the rostrum. 

Fins.—Dorsal rays 137; anal 103; pectoral 27— 
28; left pelvic 2 (right destroyed); caudal 4 + 4. 

Dorsal origin over middle of pectoral. The Ist 
through 136th dorsal rays at least 5 mm apart at 
bases; the 136th and the anterior element of the 
137th 3.8 mm apart; the two elements counted 
as the last (137th) ray 1.9 mm apart. The 135th 
and 136th rays dichotomously branched 2 or 3 
times; the two distinct elements of last ray each 
branched twice. 

Anal origin about under base of 40th dorsal 
ray. Several rays immediately preceding the 
101st are broken. The 102nd and the anterior 
element of the 103rd 4 mm apart; the two ele- 
ments counted as the last (103rd) ray 3.0 mm 
apart. Anterior of the two elements of last ray 
dichotomously branched 2 or 3 times; posterior 
branched twice. 

Pectorals originating behind posterior margin 
of opercles, subquadrate, relatively short and 
broad, without produced rays. All rays branched 
except uppermost and lowest ray of left fin and 
uppermost and two lowest of right. 

Left pelvic (right destroyed) originating 22 
mm behind anterior end of cleithrum (11 mm 
behind posterior end of maxillary, under ante- 
rior end of preopercle). Pelvic composed of 2 
rays joined for a distance of 25 mm above base. 
Outer ray 81 mm long, its free tip 56 mm long; 
part of tip of inner broken off, remaining portion 
56 mm long. At point of junction, outer ray 1.3 
mm wide; inner 2.4 mm wide. 

Caudal fin (Fig. 5) connected with dorsal and 
anal fins by the interradial membrane. Deter- 
mination of the number of caudal rays is some- 
what arbitrary. The last dorsal element articu- 
lating with a neural spine (that of the 55th caudal 
vertebra) might be regarded as a caudal ray, but 


FiGure 4. Right sagitta, horizontal diameter 12.5 mm, of 
Parabassogigas grandis, CAS 25724. Top, lateral aspect. Bot- 
tom, mesial aspect. Photographs by W. I. Follett. 


is counted as a dorsal element because of the 
small size (length 0.7 mm) of its base; the last 
anal element articulating with a haemal spine 
(that of the 55th caudal vertebra) might also be 
regarded as a caudal ray, but is counted as an 
anal element because of the small size (length 
0.9 mm) of its base. In contrast, the length of 
the base of each of the 8 caudal rays is 2.0 mm 
or more. So interpreted, the caudal fin is per- 
ceptibly differentiated from the dorsal and anal 
fins. The branching of the caudal rays (other 
than the Ist (uppermost) ray, which is dichoto- 
mously branched once) is not regularly dichot- 


394 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 17 


FIGURE 5. 


omous: the 2nd is dichotomously branched 3 to 
6 times; the 3rd, 4 or 5 times; the 4th, 4 to 6 
times; the Sth, 4 or 5 times; the 6th, 4 to 6 times; 
the 7th, 3 to 5 times; the 8th, 2 or 3 times. This 
repeated branching forms a fine fanlike pattern, 
with the rays barely distinguishable in the distal 
part of the fin. The 4 rays of each moiety of the 
caudal fin are contiguous at their enlarged bases. 
A distinct interspace separates the moieties; the 
least height of this interspace is 0.7 mm. The 4 
rays of the upper moiety articulate with the up- 
per fanlike element of the hypural complex; the 
4 rays of the lower moiety articulate with the 
similar lower hypural element. 

Lateral Line.—The lateral line of the body 
originates one orbital length above the upper end 
of the gill opening, and can be traced backward 
to a point opposite the 19th dorsal ray on the 
left side and the 15th on the right. Farther back, 
it is scarcely discernible, but it seems to descend 
to near the middle of the body height above the 
anus, behind which point it is indistinct. For 
about 110 mm behind its origin, the lateral line 
is nearly scaleless. 

On the head, the lateral line extends forward 
very faintly to a point about midway between 
the upper end of the gill opening and the anterior 
margin of the eye. From this point a rather 
prominent line of neuromasts extends vertically 
downward to about an orbit’s length above and 


Caudal fin and adjacent vertebrae of Parabassogigas grandis, CAS 25724. Print from radiograph by W. I. Follett. 


behind the upper posterior end of the maxillary, 
then downward and forward apparently to con- 
nect with a line of pores along the rostral edge. 
From the upper end of the vertical series, a faint 
line of neuromasts extends upward and forward 
a short distance, then curves downward and for- 
ward behind the eye, and then extends forward 
to behind the posterior nostril approximately 
parallel to the margin of the orbit. Then probably 
the same line extends forward in a strong up- 
ward arc to above the anterior nostril. From the 
point where the line curves downward and for- 
ward behind the eye, another line of neuromasts 
extends upward and forward about half the post- 
orbital length of the head to above the posterior 
nostril and then is abruptly deflected backward 
in line with the dorsal contour to above the ver- 
tical line on the cheek. From the end of this line, 
there seem to be a temporal commissure and 
another line extending toward the origin of the 
lateral line of the body. 

Pores.—On the edge of the snout, below the 
interval between the tubular and the crescentic 
nostrils, are two linear pores (6 mm apart), each 
4 mm long. About 3 mm above the inferior mar- 
gin of the suborbital are three smaller pores 
(about 20 mm apart). The remaining pores of the 
head are inconspicuous, except one at the angle 
of the preopercle. The anterior mandibular pore, 
a 3-mm slit with thick, scaleless margins, begins 


HUBBS & FOLLETT: OPHIDIID FISH PARABASSOGIGAS GRANDIS 395 


7: 2 
VIS es 
44: ‘egite 
Wey es 


FIGURE 6. 
by W. I. Follett. 


14 mm from the tip of the mandible and forms 
the anterior boundary of the scaled area adjoin- 
ing the symphysis. Directly behind this pore, 
and extending along the inner margin of the 
mandible, is a pore 10 mm long with thick, scale- 
less margins. Following this long pore at inter- 
vals of about 49, 32, and 33 mm are three other 
pores, at distances of about 4, 7, and 11 mm 
above the inner margin of the mandible. On the 
left side, the anteriormost of these three is the 
smallest. The narial sulcus appears to run into 
two pores internally. Near the head are several 
large pores in the lateral line, about one to a 
body segment. 

Scales.—Scales 28-—216—70/28-216-70. The 
head and body are covered with scales. Those 
of the body are cycloid, more or less oval, with 
slightly undulate margins; they are embedded, 


SAMMONS. Ay 
Ri FANG pol 
tly Vee fi 


Scale, length 6.2 mm, from near the center of the left side of Parabassogigas grandis, CAS 25724. Photograph 


not imbricated. Rows of scales, rather indistinct, 
radiate from the eye. The interorbital, preorbi- 
tals, suborbitals, opercles, preopercles, sub- 
opercles, interopercles, maxillaries, mandibles, 
and ventral aspect of the chin are densely 
scaled. A scale (Fig. 6) from near the center of 
the left side is 6.2 mm long, with eccentric (an- 
terior) focus, numerous radii in all fields, and 
fine-set circuli paralleling the scale margin; it has 
a total of 184 radii, and, in one interradial seg- 
ment of the posterior field, 112 circull. 
Rocker.—A large median bony plate (Fig. 7), 
102 mm in length and 83 mm in greatest width, 
fills most of the upper part of the anterior half 
of the body cavity. This element, known only in 
the ophidioids, has been designated the rocker 
(Rose 1961:284) or median rocker bone (Court- 
ney 1971:259). The rocker extends downward 


396 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 17 


FIGURE 7. 


Median rocker bone, length 102 mm, of Parabassogigas grandis, CAS 25724. Two of the enlarged neural spines 


are visible at the upper left. Print from radiograph by Carl B. Bowen, M.D. 


and backward from about the Sth or 6th precau- 
dal vertebra to a point directly below the 12th 
precaudal; it seems to be a process from the 6th 
precaudal. The upper surface of the plate is 
strongly calcified. The broad posterior edge is 
weakly concave; the lateral edges are strongly 
convex; the upper surface is concave, rising to 
a distinct elevation which seems to be suspend- 
ed from the dorsal wall of the body cavity. Be- 
tween the rocker and the vertebrae, and presum- 


ably supported by parapophyses, there is on 
each side a large mass of seemingly gelatinous 
tissue, obliquely truncated posteriorly; the thick 
mesentery forms the lower margin of this mass. 

Musculature.—There is a huge muscle mass 
on each side of the ventral surface of the rocker. 
The anterior part of the body cavity is largely 
filled by an immense mass of muscle extending 
forward to about opposite the end of the max- 
illary and backward to about midway between 


HUBBS & FOLLETT: OPHIDIID FISH PARABASSOGIGAS GRANDIS 397 


the isthmus and the anus. This muscle mass is 
about 130 mm wide, and is divided by a sagittal 
septum into which a high, firm keel projects 
from the rocker. This specialized musculature 
and the associated rocker presumably constitute 
a part of the sound-producing mechanism of the 
adult male (cf. Rose 1961:304; Courtney 
1971:259). 

Gas Bladder.—The gas bladder has an ex- 
tremely thick wall of fibrous tissue and fills most 
of the upper-median part of the body cavity 
above the anterior section. Anteriorly the gas 
bladder narrows to end near the posterior axil 
of the rocker. It extends back into the bifid front 
of the very thick but short kidney, the left lobe 
of which is much more massive than the right. 
Wavy light streaks on the surface of the thin- 
walled abdomen appear to have been caused by 
excessive inflation of the gas bladder—presum- 
ably as the fish rose from a great depth. There 
are remains of a rather thick fibrous mesentery 
between the kidney and the gas bladder. 

Stomach.—The stomach (which was empty) 
is a blind sac, with the pylorus behind the mid- 
dle. There are no pyloric caeca. 

Intestine.—The anterior part of the intestine 
was packed with cestodes; the posterior part 
was apparently filled with mucus. 

Testis.—The testis is a rather elongate, solid 
body full of tubules; it is extended forward about 
one-fourth the distance to the pelvic fins, and is 
incompletely fused toward the posterior end. 

Vertebrae.—Vertebrae 23 + 56 = 79. The 4th 
precaudal centrum is 15 mm in length and 26 
mm in vertical diameter; the 13th precaudal cen- 
trum is 18 mm in length and 18 mm in vertical 
diameter; the 55th caudal centrum is 7 mm in 
length and 4 mm in vertical diameter. The 4th 
and Sth precaudal centra are expanded, appar- 
ently to help support the rocker. 

Neural Spines.—The 1st neural spine is 33 
mm long, 2.5 mm in anteroposterior thickness 
at midlength, and nearly vertical. The 2nd, 3rd, 
and 4th are stout and somewhat blunt. The 2nd 
is 86 mm long and 7 mm thick; it is slightly sig- 
moid, and inclined backward at an angle of 65° 
with the axis of its centrum. The 3rd is 98 mm 
long, 6 mm thick, nearly straight, and inclined 
at 55°. The 4th is 83 mm long, 5 mm thick, slight- 
ly sigmoid, and inclined at 43°. In marked con- 
trast to these stout spines, the Sth to 12th are 
slender, with attenuate tips strongly depressed 
posteriorly. The 9th, for example, is 58 mm long, 


1.5 mm thick, and inclined at 32°. The 13th to 
16th are nearly straight and progressively longer 
and stouter. The 17th to 22nd are about 61 mm 
long, 7 mm thick, and inclined at 60°. Those be- 
hind the 22nd are progressively shorter and 
more slender and are curved backward. 

Dorsal Pterygiophores.—There are about 135 
dorsal pterygiophores. The Ist is a thin, subcir- 
cular disc about 5 mm in diameter, possibly 
fused with the 2nd. The proximal radial of the 
2nd is 70 mm long, and curves downward and 
forward (as do the 3rd to 18th); those posterior 
to the 18th are nearly straight. The Ist to 18th 
proximal radials correspond in position to the 
slender Sth to 12th neural spines. The 135th 
proximal radial is about 10 mm long. 

Haemal Spines.—The largest (2nd) haemal 
spine is 103 mm in length and 6 mm in greatest 
anteroposterior thickness; it is nearly straight 
and is inclined backward at an angle of 50° with 
the axis of its centrum. Those behind the 4th are 
progressively shorter and more slender and are 
somewhat curved backward. 

Anal Pterygiophores.—There are about 103 
anal pterygiophores. The Ist proximal radial is 
48 mm in length and 0.8 mm in anteroposterior 
thickness near its base but expanded distally to 
2.4 mm. The 2nd to 4th are 57 mm long. Six 
proximal radials precede the tip of the Ist hae- 
mal spine. The 103rd proximal radial is 18 mm 
long. 

Parapophyses.—The 9th to 22nd precaudal 
vertebrae bear expanded parapophyses; the 
largest (on the right side of the 19th vertebra) is 
56 mm in length and 7 mm in greatest width. 
These expanded parapophyses partially em- 
brace the large, thick-walled gas bladder. 

Ribs.—Measurements (in millimeters) of ribs 
on left side: 


Ist 2nd 3rd 4th Sth 6th 7th 


Length 45.0 40.0 85.0 90.0 100.0 105.0 100.0 
Width 1.5 1.0 2S 3.0 3.0 3.0 ile) 


Coloration.—When collected, the fish was 
pinkish orange (Warren E. Beadle, personal 
communication). Twenty-four hours later, a 
pinkish cast was still noticeable on the cream- 
colored head, the pale-tan sides, and the darker 
posterior region. There was a faint submarginal 
dark band along the vertical fins, and a faint dark 
bar across the middle of the pectorals. The lining 
of the mouth and gill cavity was chalky white, 


398 


becoming dusky opposite the opercles. The per- 
itoneum was brown with minute darker specks. 


NOMENCLATURE 


Parabassogigas grandis, originally described 
as Sirembo grandis, has been referred to four 
nominal genera: Sirembo Bleeker, 1858; Neo- 
bythites Goode & Bean, 1885; Bassogigas Gill 
in Goode & Bean, 1896; and Parabassogigas 
Nybelin, 1957. (See Synonymy.) 

We follow Nybelin (1957:298) in referring Si- 
rembo grandis to the nominal genus Parabas- 
sogigas, of which grandis is the type-species by 
original designation. 

In referring Parabassogigas grandis to the 
family Ophidiidae, we follow Cohen and Nielsen 
(in press). 


ACKNOWLEDGMENTS 


We thank Richard S. Croker, formerly of the 
California Department of Fish and Game, and 
Willard R. Yuna, Fish and Game Warden, for 
enabling us to obtain the California specimen of 
Parabassogigas grandis; Warren E. Beadle of 
Walteria, California, and R. G. Hamilton of 
Venice, California, for saving the floating fish 
and for presenting it to the California Academy 
of Sciences; Ethelwynn Trewavas of the British 
Museum (Natural History) for data on the ho- 
lotype of Sirembo grandis; Tokiharu Abe of the 
University of Tokyo for data on a large Japanese 
specimen of Parabassogigas grandis; John E. 
Fitch of the California Department of Fish and 
Game for diagnosis of a sagitta of the California 
specimen; Jean-Claude Hureau of the Muséum 
National d’Histoire Naturelle, Paris, for con- 
firming our identification of the California spec- 
imen; Lillian J. Dempster of the California 
Academy of Sciences for extensive assistance 
with the manuscript; and Maurice C. Giles of 
the California Academy of Sciences for expertly 
enlarging the photographs and printing the ra- 
diographs. 


LITERATURE CITED 


BLEEKER, PIETER. 1858. Vierde bijdrage tot de kennis der 
ichthyologische fauna van Japan. Acta Soc. Sci. Indo- 
Neerl. 3: 1-46. 

BRAUER, AUGUST. 1906. Die Tiefsee-Fische. Wiss. Ergeb. 
Dtsch. Tiefsee-Exped. ‘* Valdivia’? 1898-1899, 15(1):1—432, 
pls. 1-16, figs. 1-176. 

COHEN, DANIEL M. 1963. The publication dates of Goode 
and Bean’s Oceanic Ichthyology. J. Soc. Bibliogr. Nat. 
Hist. 4(3): 162-166. 


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COURTNEY, WALTER R., JR. 1971. Sexual dimorphism of the 
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DoLLo, Louis. 1906. Neobythites Brucei, poisson abyssal 
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EAGLE, RODNEY J. 1969. First records from the northeastern 
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FoLLeTtT, W. I. 1970. Benthic fishes cast ashore by giant 
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GARMAN, SAMUEL. 1899. Reports on an exploration off the 
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GooDE, GEORGE BROWN, AND TARLETON H. BEAN. 1885. 
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GreY, MARION. 1956. The distribution of fishes found below 
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GUNTHER, ALBERT. 1877. Preliminary notes on new fishes 
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1887. Report on the deep-sea fishes collected by 
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Husss, Cart L. 1919. A comparative study of the bones 
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HUBBS & FOLLETT: OPHIDIID FISH PARABASSOGIGAS GRANDIS 399 


NorMAN, J. R. 1939. Fishes. Sci. Rep. John Murray Exped., 
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. 1954. Sur la distribution géographique et bathymét- 

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. 1957. Deep-sea bottom-fishes. Rep. Swed. Deep-sea 
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TANAKA, SHIGEHO. 1908. Notes on some rare fishes of Ja- 
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THOMSON, C. WYVILLE. 1880. Appendix to the introduction 
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Vol. XLI, No. 18, pp. 401-426; 14 figs.; 6 tables 


it 


it 


Pon es 


ae 


NH 


x 


PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


TWO NEW AUSTRALIAN VELVETFISHES, GENUS 
PARAPLOACTIS (SCORPAENIFORMES: APLOACTINIDAE), 
WITH A REVISION OF THE GENUS AND COMMENTS ON 

THE GENERA AND SPECIES OF THE APLOACTINIDAE 


By 
Stuart G. Poss 
University of Michigan Museum of Zoology, Ann Arbor, Michigan 48109 


and 
William N. Eschmeyer 
California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 


ABSTRACT: The velvetfish genus Paraploactis, formerly consisting of a single species, P. trachyderma 
Bleeker, 1865, from Queensland is revised to include several species previously referred to Cocotropus and 
two new species: P. intonsa and P. pulvinus from Western Australia. Specimens referred to four nominal 
species: Tetraroge kagoshimensis Ishikawa, 1904, in Jordan and Starks from Japan, Cocotropus obbesi 
Weber, 1913, from the Sulu Sea, Aniculerosa taprobanensis Whitley, 1933, from Sri Lanka, and Cocotropus 
hongkongensis from Hong Kong make up an unresolved complex also referable to Paraploactis. Members 
of the Paraploactis kagoshimensis complex are the least specialized of the species of Paraploactis, P. 
trachyderma and P. pulvinus being the most specialized. This specialization is reflected in increases in 
depression of the body, development of cirri on the snout, fleshiness of the ventral surface of the mandible, 
and development of a unique fleshy pad on the isthmus. Scanning electron photomicrographs show that 
scales of species of Paraploactis form spinous points. The species are described and figured, phenetic and 
cladistic analyses presented, and a key provided. Tables of the nominal genera and species of the Aploac- 
tinidae are given as is a discussion of features characterizing the family. 


December 28, 1978 


INTRODUCTION 


Among the families of the Scorpaenoidei (sen- 
su Greenwood et al. 1966), perhaps none is more 
unusual than the Aploactinidae. These fishes, 
about 24 species in number, are characterized 
by having a dorsal fin originating far forward on 
the cranium, a head invested with knoblike 
lumps, and a body usually covered with modi- 
fied, prickly scales (hence, the name velvetfish- 
es). In addition to these features, and quite un- 


usual to scorpionfishes and their relatives, 
aploactinids show a loss of pungent fin spines, 
and some may be nonvenomous. If so, they 
would seem even more remarkable as they ap- 
pear closely related to the notoriously venomous 
synanceline and tetrarogine scorpaenids. 
Essentially, all that is known of these fishes 
are a single review (Whitley 1933) based on Aus- 
tralian forms and relying heavily on the litera- 
ture, a detailed regional study of the taxonomy 


402 


and morphology of three Japanese species (Mat- 
subara 1943), a small body of information in 
original descriptions, and a few faunal lists. 
Aploactinids occur at scattered localities 
throughout the western Pacific and Indian 
oceans. They appear to be more numerous in 
the Indonesian and Australian regions, but this 
may reflect an inadequate knowledge about oth- 
er areas. Members of the group are presumed to 
be primarily restricted to continental margins, 
though a few insular records are known: Waite 
1903 (Lord Howe); Seale 1935, and Fowler 1959 
(Fiji); Eschmeyer and Poss 1976 (Raoul). Little 
if any information exists on the depths at which 
some species have been taken. As a group they 
are known to occur from nearshore to 510 m. 
Based on limited information for a few species, 
it is suspected that most live within interstices 
on rocky or coral-rubble bottoms, bottoms with 
coralline algae, or among vegetation. Their vir- 
tual absence in museum collections (most being 
known from one or a few individuals) can pre- 
sumably be explained by their presence in these 
difficult-to-sample habitats. They are small in 
size and cryptic in appearance, factors which 
would further add to the difficulty of their cap- 
ture. 

The spelling Aploactidae has commonly been 
used in the literature (e.g. Waite 1921, 1923; Jor- 
dan 1923; McCulloch 1929; Whitley 1933; Berg 
1940; Scott 1942; Munro 1955, 1967; Scott 1962; 
McKay 1970; Scott et al. 1974). This compound 
name was derived by Temminck and Schlegel 
(1843:51) from the Greek aplos (simple) + aktis 
(ray), in reference to the unbranched fin rays in 
the Japanese species Aploactis aspera. The 
spelling Aploactinidae, though less common 
(Greenwood et al. 1966; Nelson 1976; Scott 
1976), is grammatically correct. However, ac- 
cording to amended Article 29 of the Interna- 
tional Code of Zoological Nomenclature (In- 
ternational Commission on Zoological 
Nomenclature 1974:81), a family group pro- 
posed before 1961 based on an incorrectly 
formed stem is not to be amended for that reason 
if it is in general current use. (General current 
use is defined (ibid. 1964:87, 88, Article 79) as 
requiring at least five different authors in at least 
ten publications.) Thus, the name Aploactidae, 
based on an incorrectly formed stem, can be 
considered the correct spelling under the rule in 
effect at this writing. However, the Editorial 
Committee of the Commission has recently pro- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


posed (ibid. 1977:170) removal of the provision 
(Article 29d) preserving original spelling. We 
suggest this amendment will pass and prefer the 
corrected spelling Aploactinidae for this family. 

Although Matsubara (1943) in his classical 
work on Japanese scorpaenoids considered the 
group a subfamily (Aploactinae) of the broad 
family Scorpaenidae and demonstrated its rela- 
tionship to other scorpaenoids, we follow most 
authors in recognizing a family Aploactinidae. 
We include in the Aploactinidae species referred 
to the family Bathyaploactidae (Whitley 1933), 
which were not treated by Matsubara (1943). 

We discuss the features which have been used 
to characterize the family and present a list of 
all the nominal species and their current status. 
This is accomplished in a section preceding our 
review of the genus Paraploactis. 

The genus Paraploactis includes the largest 
aploactinids. Several of the Australian species 
are fairly well represented in museum collec- 
tions. Specimens from other areas are rare and 
present problems in classification at the species 
level, as is shown in the text. 

Among fishes of the genus Paraploactis, the 
isthmus may bear a fleshy pad. This structure in 
its most developed condition might be used as 
a sucking disk. We describe two new Australian 
species, review and more precisely define the 
limits of Paraploactis, cite new records, and 
speculate, where possible, on the biology and 
phyletic relationships of the species. 


METHODS 


Methods for taking fin ray counts and mea- 
surements follow those of Eschmeyer (1969). 
The last two rays of the dorsal and anal fin are 
borne on a single pterygiophore and counted as 
one ray. Spines and rays are difficult to distin- 
guish and all counts were checked against radio- 
graphs. Several nonstandard counts and mea- 
surements useful in differentiating species of 
Paraploactis were made: papillae per mandibu- 
lar cirrus (fleshy projections counted on fourth 
left cirrus); number of cirri along outer lower 
margin of mandible (counted on both sides of 
mandible); number of cirri along inner lower 
margin of mandible (counted on both sides of 
mandible); length of cirri on snout (length of first 
cirrus anterior to left lachrymal bone); length of 
cirri on nasals (length of longest cirrus on left 
nasal bone); distance between interorbital ridges 
(from outer edges of interorbital ridges at pos- 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 


terior border of interorbital pit); pad length 
(maximum length of pad); pad width (maximum 
width of pad). 

Photomicrographs of the scales were made 
with a JEOL SJM—U3 scanning electron micro- 
scope at the Scanning Electron Microscope— 
Electron Microprobe Analyzer Laboratory at 
the University of Michigan (SEM—EMPA Lab). 
The skin was glow discharge coated with a 250- 
300 A layer of gold prior to observation. 

All numerical computations were performed 
with the MTS—MIDAS algorithms developed by 
the Statistical Research Laboratory at the Uni- 
versity of Michigan. 

Abbreviations of depositories of specimens 
are: AMS—dAustralian Museum, Sydney; 
ANSP—Academy of Natural Sciences of Phil- 
adelphia; BMNH—British Museum (Natural 
History), London; CAS—California Academy 
of Sciences, San Francisco; CMFRI—Central 
Marine Fisheries Research Institute, Mandapam 
Camp, India; CSIRO—Commonwealth Scientif- 
ic Industrial Research Organization, Cronulla, 
Australia; FRSKU—Fisheries Research Sta- 
tion, Kyoto University, Kyoto, Japan; MCZ— 
Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge; QM—Queensland Museum, 
Brisbane, Australia; RMNH—Rijksmuseum van 
Natuurlijke Historie, Leiden; SAM—South 
Australian Museum, Adelaide, Australia; SU— 
Stanford University Collection (at CAS); 
USNM—United States National Museum of 
Natural History, Washington; WAM—Western 
Australian Museum, Perth; UW—University of 
Washington College of Fisheries, Seattle; 
ZMA—Instituut voor Taxonomische Zoologie, 
Zoologische Museum, Amsterdam. 


ACKNOWLEDGMENTS 


The authors are indebted to the curators and 
staffs of a number of museums for making spec- 
imens available. Without their help this study 
would not have been possible. We express our 
sincere thanks to Drs. Gerald Allen (WAM), 
James Bohlke (ANSP), John Glover (SAM), 
Phillip Heemstra (CSIRO), Karel Liem (MCZ), 
R. J. McKay (QM), Ian Munro (CSIRO), Izumi 
Nakamura (FRSKU), Han Nijssen (ZMA), John 
Paxton (AMS), E. G. Silas (CMFRI), Victor 
Springer (USNM), and Arthur Welander (UW) 
for loaning material to us. Special thanks are due 
M. Boeseman (RMNH) and Alwyne Wheeler 
(BMNH) for their considerable assistance in ex- 


403 


amining types and in making material available. 
We thank M. L. Connoly (BMNH), Karsten 
Hartel (MCZ), Susan Karnella (USNM), and G. 
Venkataraman (CMFRI) for their assistance in 
sending specimens. Ellie Baker (UMMZ) assist- 
ed in the shipment of specimens and in preparing 
radiographs. Others at the University of Michi- 
gan who provided assistance were Dr. Reeve 
Bailey, George Estabrook, Dolores Kingston, 
Dr. Robert R. Miller, Ken Rose, and Dr. Gerald 
Smith. Peggie Hollingsworth (SEM—-EMPA 
Lab) gave technical assistance during use of the 
scanning electron microscope. Computer funds 
were made available by the UMMZ. Acknowl- 
edgment is due Dr. Bruce Collette (USNM), 
Karsten Hartel, and Santo Vitale (William K. 
Vanderbilt Museum) for their help in the search 
for the holotype of Erosa australiensis , Dr. Ter- 
uya Uyeno, Dr. Ryoichi Arai, and Mr. Katsu- 
suke Meguro for their help in search of the ho- 
lotype of Tetraroge kagoshimensis, and Dr. 
Kaza V. Rama Rao and Alwyne Wheeler for 
help in search of the holotype of Aniculerosa 
taprobanensis. The drawings in this study were 
made by Holly Wong. William Ruark (CAS) pre- 
pared radiographs. Melissa Barbour, Lillian 
Dempster, W. I. Follett, Dr. Warren Freihofer, 
James Gordon, Howard Hammann, Karren 
Hakanson, Dr. Tomio Iwamoto, Ken Lucas, 
Betty Powell, Kathy Smith, and Pearl Sonoda, 
all of CAS, assisted throughout the study. 


GENERA AND SPECIES OF THE FAMILY 
APLOACTINIDAE 


In the only revision of the group, Whitley 
(1933) characterizes the Aploactinidae by their 
having 1) head spines which are not pungent but 
developed as knoblike processes, 2) modified 
scales which form spinous points or prickles, 3) 
anal spines which are either indistinct or absent 
entirely, 4) a dorsal fin which originates on the 
cranium, 5) fewer than four pelvic rays, the in- 
nermost of which is not adnate to the body, 6) 
no palatine teeth, 7) no gill slit behind the last 
arch, 8) no free pectoral rays, and 9) a toothed 
vomer. Of these features the last two are found 
in the vast majority of fishes and probably rep- 
resent features primitive to the group (and dis- 
regarding the fact that some individuals of Par- 
aploactis species lack vomerine teeth). Features 
4, 5, 6, and 7, while apparently derived, are also 
present in the closely related tetrarogine and 
synanceline scorpaenids. Features 1, 2, and 3 


404 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


TABLE 1. THE NOMINAL GENERA OF APLOACTINID FISHES AND THEIR CURRENT STATUS (TYPE-SPECIES IN PARENTHESES). 


Genus (type-species) 


Acanthosphex Fowler, 1938 (leurynnis) 
Adventor Whitley, 1952 (elongatus) 
Aniculerosa Whitley, 1933 (taprobanensis) 
Aploactis Temminck & Schlegel, 1843, in 
Siebold, (aspera) 
Aploactoides Fowler, 1938 (philippinus) 
Aploactisoma Castelnau, 1872 (schomburgki) 
Bathyaploactis Whitley, 1933 (curtisensis) 
Cocotropus Kaup, 1858 (echinatus) 
Erisphex Jordan & Starks, 1904 (potti) 
Insopiscis Whitley, 1933 (altipinnis) 
Kanekonia Tanaka, 1915 (florida) 
Karumba Whitley, 1966 (ornatissima) 
Kleiwegia de Beaufort, 1952 (dezwaani) 
Membracidichthys Whitley, 1933 (obbesi) 
Neoaploactis Eschmeyer & Allen, in press 
(tridorsalis ) 
Paraploactis Bleeker, 1865 (trachyderma) 
Peristrominous Whitley, 1952 (dolosus) 
Sthenopus Richardson, 1848 (mollis) 
Trichopleura Kaup, 1858 (mollis) 


Status 


Remarks 


Acanthosphex 
Adventor 
Paraploactis 


Aploactis 
Erisphex 
Aploactisoma 
Bathyaploactis 
Cocotropus 
Erisphex 
Cocotropus (?) 
Kanekonia 
Karumba 
Acanthosphex 
Paraploactis 


Neoaploactis 
Paraploactis 
Peristrominous 
Sthenopus 
Stenopus 


See original description 
Whitley 1952a 
This paper 


See Matsubara 1943 
See Matsubara 1943 
See Whitley 1933 

See original description 
This paper 

See Matsubara 1943 
This paper 

See Tanaka 1918 

See original description 
See Weber & de Beaufort 1962 
This paper 


See original description 

This paper 

See Whitley 1952a 

See Weber & de Beaufort 1962 
See Weber & de Beaufort 1962 


are specializations which are found only among 
species we include in the Aploactinidae. Not all 
species so included, however, possess all three 
features. Some, such as Erisphex potti and Ka- 
nekonia aniara, have pungent head spines. Oth- 
ers, such as Kanekonia florida and Sthenopus 
mollis , lack all or nearly all prickly scales on the 
skin and have instead a smooth, fleshy skin. 
Several species within the genera Erisphex, Ka- 
nekonia, and Cocotropus have pungent anal 
spines. These spines tend to be weakly devel- 
oped in large specimens and usually involve only 
the first spine. These genera are generally re- 
garded as intermediate between the remainder 
of the aploactinid genera and those of scorpaen- 
id subfamilies (see for example, Jordan and 
Starks 1904; Whitley 1933; Matsubara 1943; 
Smith 1958). 

Species of the genera Acanthosphex and 
Bathyaploactis have somewhat pungent spines. 
Both genera are characterized by having the gill 
slits reduced to small openings, an apparently 
specialized feature which sets them apart from 
other aploactinids. They also have large preor- 
bital spines, and fewer velvety prickles on the 
skin. 

Although limited to but three Japanese spe- 
cies, the detailed anatomical work of Matsubara 
(1943) provides several additional and apparent- 
ly specialized features which further character- 
ize the Aploactinidae. His osteological exami- 


nation of Erisphex pottii, Paraploactis 
kagoshimensis, and Cocotropus masudai re- 
vealed these forms to have: 1) six as opposed to 
seven branchiostegals (see below); 2) an appar- 
ently unique configuration and modification of 
the urohyal (see also Kusaka 1974); 3) an epiot- 
ic-opisthotic articulation in addition to the more 
common articulation of the epiotic with the ex- 
occipital, supraoccipital, pterotic, and parietal; 
and 4) the lateral ethmoids (ectethmoids) widely 
separated anteriorly from the vomer by carti- 
lage. The last feature is known elsewhere among 
scorpaenoids in fishes of the scorpionfish 
subfamily Pteroinae. The first three features are 
not known to occur in other fishes of the sub- 
order Scorpaenoidea. Members of the Tetraro- 
ginae lack an epiotic-opisthotic articulation. The 
position of the anterior neural spines relative to 
the cranium is different in tetrarogines and 
aploactinids. The aploactinids Peristrominous 
dolosus and Adventor elongatus do not have the 
anterior dorsal spines originating on the crani- 
um. This suggests that the advancement of dor- 
sal spines onto the cranium may have taken 
place more than once in these fishes. 
Examination of a cleared and alizarin-stained 
specimen of Paraploactis intonsa (CAS 39600) 
revealed the presence of seven (both right and 
left sides) rather than six branchiostegal rays. 
McAllister (1968) states that five or six bran- 
chiostegal rays are present in one (unspecified) 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 


TABLE 2. 


Original name (Species, genus) 


achrurus, Erisphex, Regan, 1905 
altipinnis, Cocotropus, Waite, 1903 
aniara, Kanekonia, Thompson, 1967 


aspera, Synanceia (Aploactis), Richardson, 1844 

asperrimus, Prosopodasys, Gunther, 1860 

australiensis, Erosa, Borodin, 1932 

curtisensis, Bathyaploactis, Whitley, 1933 

dermacanthus, Apistus, Bleeker, 1852 

de Zwaani, Cocotropus, Weber & de Beaufort, 1915 

dolosus, Peristrominous, Whitley, 1952a 

echinatus, Corythobatus, Cantor, 1850 

elongatus, Membracidichthys (Adventor), Whitley, 1952 

florida, Kanekonia, Tanaka, 1915 

hongkongiensis, Cocotropus, Chan, 1966 

horrenda, Aploactisoma milesii, Whitley, 1933 

intonsa, Paraploactis, Poss & Eschmeyer n. sp. 

kagoshimensis, Cocotropus, Ishikawa, 1904, in 
Jordan & Starks 

lichen, Aploactis, de Vis, 1884 

leurynnis, Acanthosphex, Jordan and Seale, 1906 

masudai, Cocotropus, Matsubara, 1943 

milesii, Aploactis, Richardson, 1850 

mollis, Sthenopus, Richardson, 1848 

monacanthus, Tetraroge, Gilchrist, 1906 

obbesi, Coccotropus , Weber, 1913 

ornatissimus, Bathyaploactis curtisensis, Whitley, 1933 

philippinus, Aploactoides, Fowler, 1938 

pottii, Cocotropus, Steindachner, 1896 

queenslandica, Kanekonia, Whitley, 1952 

roseus, Cocotropus (Tetraroge), Day, 1875 

schomburgki, Aploactisoma, Castelnau, 1872 

sieboldi, Aploactus, Kaup, 1852 

steinitzi, Cocotropus, Eschmeyer & Dor, in press 

taprobanensis, Aniculerosa, Whitley, 1933 

tridorsalis, Neoaploactis, Eschmeyer & Allen, in press 

trachyderma, Paraploactis, Bleeker, 1865 


405 


THE NOMINAL SPECIES OF APLOACTINID FISHES AND THEIR CURRENT STATUS. 


Status 


Remarks or Identification 
Source 


Erisphex potti 
Cocotropus altipinnis 
Kanekonia aniara 


Aploactis aspera 
Uncertain 

Paraploactis trachyderma 
Bathyaploactis curtisensis 
Cocotropus dermacanthus 
Acanthosphex leurynnis 
Peristrominous dolosus 
Cocotropus echinatus 
Adventor elongatus 
Kanekonia florida 
Paraploactis hongkonginesis 
Uncertain 

Paraploactis intonsa 


Paraploactis kagoshimensis 
Paraploactis trachyderma 
Acanthosphex leurynnis 
Cocotropus masudai 
Aploactisoma milesii 
Sthenopus mollis 
Cocotropus monacanthus 
Paraploactis obbesi 
Karumba ornatissima 
Erisphex philippinus 
Erisphex pottii 

Kanekonia queenslandica 
Cocotropus roseus 
Aploactisoma milesii 
Aploactis aspera 
Cocotropus steinitzi 
Paraploactis kagoshimensis (?) 
Neoaploactis tridorsalis 
Paraploactis trachyderma 


See Matsubara 1943 

This paper 

See original description; 
generic placement uncertain 

See Matsubara 1943 

See original description 

This paper 

See original description 

See Herre 1951 

See Weber & de Beaufort 1962 

See Whitley 1952a 

See original description 

See Whitley 1952a 

See Tanaka 1918 

This paper 

See Scott 1976 

This paper 


This paper 

This paper 

See Ramaiyan & Rao 1970 
See original description 
Whitley 1933 

See Weber & de Beaufort 1962 
See Smith 1958 

This paper 

See Whitley 1966 

See original description 
See Matsubara 1943 

See Whitley 1952b 

This paper 

See Whitley 1933 

See Matsubara 1943 

See original description 
This paper 

See original description 
This paper 


aploactinid genus examined by him. In P. inton- 
sa the anteriormost branchiostegal ray is little 
more than a small sliver of bone which is weakly 
attached to the ceratohyal. 

Until the features noted by Matsubara for 
three species of Japanese aploactinids can be 
examined in the remainder of the group, the pre- 
cise limits of the family and its status as an entity 
distinct from the related syanceiine and tetrar- 
ogine scorpaenids will remain obscure. We ten- 
tatively include the nominal genera listed in Ta- 
ble 1 and the nominal species listed in Table 2 
in the family Aploactinidae. 


Genus Paraploactis Bleeker 


Paraploactis BLEEKER, 1865:168-169 (type-species Para- 
ploactis trachyderma Bleeker, 1865, by original designa- 
tion, monotypic). BLEEKER 1876a:200 (description). BLEEK- 


ER 1876b:8 (description). WHITLEY 1933:100 (description). 
MARSHALL 1965:420, 434 (in key; compiled). ScoTT 
1976:207—208 (compared with Aploactisoma). 

Membracidichthys WHITLEY, 1933:102 (type-species Cocco- 
tropus [sic] obbesi Weber, 1931, by original designation). 

Aniculerosa WHITLEY, 1933:101 (type-species Aniculerosa 
taprobanensis Whitley, 1933, by original designation, 
monotypic). 


DiaGNosis.—Aploactinid fishes densely cov- 
ered with scales modified to form prickles, an 
elaborately sculptured head, and in some spe- 
cies a fleshy pad on isthmus. Frontal bone with 
two prominent ridges that diverge posteriorly, 
forming a pyriform depression between them. 
Dorsal fin originating on cranium over middle to 
rear of orbit. Dorsal fin membrane between 
spines 3 and 4 usually deeply incised so that first 
four spines nearly form a separate fin. Dorsal 


406 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


100 


FIGURE 1. 


20 


20 


160 


Distribution map for known records of species of Paraploactis. Circles represent P. trachyderma; circles with 


star, P. pulvinus; triangle, P. intonsa; squares, members of P. kagoshimensis complex. Not shown are two records of specimens 
belonging to P. kagoshimensis complex (P. taprobanensis) from Sri Lanka. 


and anal fin spines often flexible and difficult to 
distinguish from soft rays; all fin rays un- 
branched. Total dorsal fin elements 22-24; with 
13-15 spines and 8-10 soft rays. Anal fin with 
1 spine (rarely 2) and 7-10 soft rays; total ele- 
ments 8-11. Pectoral fin rays 13-14, usually 14, 
all unbranched. Pelvic rays I + 3. Head and 
body densely covered with prickly scales, larger 
scales above lateral line behind head. Lateral 
line scales, particularly anteriorly, as prominent 
knobs, total 11-15. Minute villiform teeth in 
jaws; palatine teeth absent; vomerine teeth 
sometimes absent or reduced. Gill rakers as 
blunt knobs, 6-11 total; 0-1 on upper arch, 5—9 
on lower arch. Head with blunt, obtuse spines. 
Lachrymal bone (infraorbital 1) movable and 


can be ‘locked’ outward; three spines on lach- 
rymal bone, anteriormost small and pointing 
ventrally, posterior two large, one above the 
other, pointing mostly to rear with mouth 
closed. Suborbital portion of infraorbital series 
as a strong ridge with a lump in middle (on in- 
fraorbital bone 2), followed by two or three con- 
spicuous obtuse spines (on infraorbital bone 3). 
Postorbital portion of infraorbital series with a 
single ossified element. Preopercle with five 
spines, uppermost largest, with a supplemental 
spine or lump at its base. No slit behind last gill 
arch. Gasbladder absent. Gasbladder muscle 
present, originating on occipital part of cranium, 
attaching to dorsal portion of cleithrum, passing 
posteriorly ventral to pleural ribs, and inserting 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 407 


FIGURE 2. 
right: P. intonsa (holotype). Lower left: P. trachyderma (AMS 1.12514). Lower right: P. pulvinus (holotype). 


via distinct tendons to paraphophyses 6—11 (two 
species examined). Vertebrae 26-28. Branchio- 
stegals 6 or 7. 

DISCUSSION AND RELATIONSHIPS.—Many 
aploactinid species are in monotypic genera. 
Cocotropus, on the other hand, has been a 
‘catchbasket’ genus in which a number of spe- 
cies have been described and to which other 
species have been assigned, including nominal 
species which we refer to Paraploactis. While 
limited material precludes definitive statements, 
we believe Cocotropus to be restricted to the 
following species: C. echinatus (Cantor, 1849; 
the type-species of Cocotropus Kaup, 1858), C. 
dermacanthus (Bleeker, 1852), C. masudai Mat- 
subara, 1943, C. monocanthus (Gilchrist, 1906), 
C. roseus (Day, 1875), C. altipinnis Waite, 1903, 
and C. steinitzi Eschmeyer and Dor in press. We 


Frontal view of species of Paraploactis. Upper left: member of P. kagoshimensis complex (CSIRO 3237). Upper 


refer Tetraroge kagoshimensis and its possible 
synonyms (most of which have been described 
in Cocotropus or referred to it) to Paraploactis 
as the ‘‘P. kagoshimensis complex.’’ The genus 
Paraploactis otherwise contains P. trachyderma 
and the two new species described below. The 
distribution of the species of the genus Para- 
ploactis is shown in Figure 1. 

On the basis of overall similarity, species of 
Paraploactis are most closely related to species 
of the genus Cocotropus and may be considered 
a sister-group of all or part of that genus. Co- 
cotropus monocanthus, in particular, is very 
similar in appearance to species of Paraploactis. 
Species of Paraploactis are, however, more spe- 
cialized in a number of features. They have the 
posterior margins of the interorbital ridges di- 
verging posteriorly (Fig. 2); the interorbital 


408 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


FIGURE 3. 
P. intonsa (holotype). C. P. trachyderma (AMS 1.12514). D. P. pulvinus (holotype). (Drawn by Holly Wong.) 


ridges of nearly all aploactinids and their rela- 
tives which possess them extend directly back 
without diverging posteriorly. Species of Para- 
ploactis differ from those of Cocotropus in hav- 
ing the membrane of the dorsal fin between 
spines 3 and 4 deeply incised so as to form a 
nearly separate fin. The degree of incision is less 
in representatives of the P. kagoshimensis com- 
plex and greater in other Paraploactis species. 
Species of Paraploactis also have more de- 
pressed bodies than those of Cocotropus. 

The fleshy pad on the isthmus and the asso- 
ciated modification of the underside of the head 
in some species of the genus Paraploactis rep- 
resent a specialization unique among fishes. An 


Ventral view of head in species of Paraploactis. A. Member of P. kagoshimensis complex (AMS IA.3951). B. 


increase in complexity, as reflected in changes 
in the size and shape of the pad and apparently 
concomitant modifications of the external sur- 
face of the ventral part of the head is found with- 
in the genus. Representatives of the Paraploac- 
tis kagoshimensis complex show no development 
of a pad on the isthmus and show a full comple- 
ment of cirri arranged in an inner and outer row 
on the ventral surface of the mandible (Fig. 3A). 
These fishes possess a somewhat fleshy projec- 
tion of the anteriormost portion of the isthmus. 
A similar fleshy extension of the isthmus is pres- 
ent in all other aploactinids and, interestingly, 
in Ptarmus gallus, pataecids, and Congiopodus 
leucomentopon. It is reasonable to speculate 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 409 


FIGURE 4. 


Scanning electron micrographs of scale from upper back of species of Paraploactis. Upper left: member of P. 


kagoshimensis complex, 185x (AMS E.2944). Upper right: P. intonsa 165x (WAM 5378). Lower left: P. trachyderma, 120x 


(AMS 1.12514). Lower right: P. pulvinus , 340x (WAM P23735). 


that such a condition preceded the formation of 
the fleshy pad on the isthmus seen in other spe- 
cies of Paraploactis. 

In Paraploactis trachyderma and Paraploac- 
tis intonsa a pad about 4—'4 as wide as long is 
present on the isthmus. It is formed of thick- 
ened, somewhat spongy connective tissue and 
is less fleshy in P. intonsa (Fig. 3B) than in P. 
trachyderma (Fig. 3C). In the latter the ventral 
surface of the mandible is quite fleshy, the cirri 
along its inner margin are absent or appear as 


small fleshy stubs, and the cirri along its outer 
margin are thickened and show no or a few well- 
developed papillae on the surface of the cirri. 

Paraploactis pulvinus shows the greatest de- 
velopment of the pad (Fig. 3D). The pad is about 
¥2—-34 as wide as long and very fleshy. The ven- 
tral surface of the mandible is also extremely 
fleshy, cirri are entirely absent along its inner 
margin, and those along the outer margin are 
fleshy and flattened as in P. trachyderma, but to 
an even greater extent. 


410 


kagoshimensis intonsa trachyderma pulvinus 
complex 
Ficure 5. Cladogram representing the presumed phylo- 


gentic relationships among species of Paraploactis (see text 
for discussion). 


The adaptive or mechanical significance of the 
pad can as yet only be surmised. To our knowl- 
edge no aquarium or field observations have 
been made. The apparent trend toward loss of 
pigment and cirri on the ventral surface of the 
mandible and pad, and the shape and location 
of the pad suggest an intimate contact between 
the pad and the substrate. The pad may have 
originated as a structure to enhance stability on 
the bottom. With an increase in size and flesh- 
iness, and an association with the overlying 
musculature, the pad might function as part of 
a sucking disc. The pad itself consists only of 
connective tissue. It is unknown if these fishes 
live in strong surge areas as some species of 
other fish groups (cyclopterids, gobiesocids, go- 
bioids, sisorids) that have evolved ventral ad- 
hesive organs. 

SEM photomicrographs of the prickly scales 
of species of Paraploactis are presented in Fig- 
ure 4. Each scale consists of a circular base sup- 
porting a spinous process which emerges lat- 
erally and slightly posteriorly from the center of 
the base. The scale is covered with cornified ep- 
idermis, or cuticle, with numerous secondary 
projections. There is variation in prickle size and 
shape over the body. They are larger and tend 
to be bladelike on the upper back and smaller 
and more conical ventrally and posteriorly. 

The presumed cladistic relationships among 
species of the genus Paraploactis are presented 
in Figure 5. This is based primarily on the de- 
velopment of the pad, including the size of the 
pad, the loss of cirri and pigment on the ventral 
surface of the mandible, and an increase in flesh- 
iness of both the pad and the underside of the 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


head. That P. pulvinus is the most specialized, 
is further evidenced by the heavily ossified and 
sculptured interorbit and the large fleshy cirri on 
the snout. Although the pad is an unusual and, 
in fact, unique feature, we do not believe that 
its presence warrants generic separation of those 
having the pad from those lacking the pad. 

PRINCIPAL COMPONENTS ANALYSIS.—A 
45 x 46 matrix representing 45 characters for 46 
individuals was standardized and the first six 
principal components extracted. The variables 
scored are given in Tables 4-7. All measure- 
ments were divided by standard length prior to 
standardization of the character variables (stan- 
dard length was not otherwise included in the 
analysis). As one individual (CSIRO C2429) had 
a markedly abnormal anal fin, variables describ- 
ing anal fin length and anal rays were excluded 
in forming the matrix. The specimen was also 
removed from an analysis using all 45 charac- 
ters, the resultant scatter plots being only slight- 
ly different. Five specimens of P. trachyderma 
for which data were unavailable for all variables 
were also excluded. Results for the first two 
components are summarized in Figure 6. 

Principal Component I. The major axis of 
variation in the sample has an eigenvalue of 10.8 
and accounts for 24% of the total variance. 
Characters with the highest factor loadings on 
this axis are: body depth (.26), pad width (.25), 
pad length (.25), and snout to base of second 
dorsal spine (.25). 

Principal Component II. This factor has an 
eigenvalue of 5.4 and accounts for 12% of the 
total variance. Characters with the highest fac- 
tor loadings of this axis are: length of first dorsal 
spine (.37), length of third dorsal spine (.35), 
length of second dorsal spine (.33), length pen- 
ultimate dorsal spine (.31), and length of last 
dorsal spine (.28). 

Principal Component III. This factor has an 
eigenvalue of 4.2 and accounts for 9% of the 
total variance. Characters with the highest fac- 
tor loadings on this axis are: vertebrae (.39), 
dorsal rays (.33), cirri on inner margin of lower 
jaw (.28), and number of papillae per cirrus (.28) 
(also anal rays (.38) and anal fin length (.29) 
when included). 

Principal Component IV. This factor has an 
eigenvalue of 2.9 and accounts for 6% of the 
total variance. Length of cirri on snout showed 
a loading of .27. Characters with notable nega- 
tive loadings on this axis are: caudal fin length 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 


(—.34), jaw length (—.28), snout to base of third 
dorsal spine (—.26), and snout to base of fourth 
dorsal spine (—.25). 

Principal Component V. This factor has an 
eigenvalue of 2.2 and accounts for 5% of the 
total variance. Characters with the highest load- 
ings on this axis are: left gill rakers (.35), right 
gill rakers (.34), and orbit diameter (.27). Char- 
acters with negative loadings on this axis are: 
pectoral fin length (—.34) and interorbit (—.26). 

Principal Component VI. This factor has an 
eigenvalue of 2.0 and accounts for 5% of the 
variance. The highest positive loading on this 
axis is fifth dorsal spine length (.23). Notable 
negative loadings on this axis are: distance be- 
tween interorbital ridges (—.44), left gill rakers 
(—.39), right gill rakers (—.34), and last dorsal 
spine (—.30). 

It is thought that the first three axes summa- 
rize the major differences among species of Par- 
aploactis. Principal components I and II suggest 
the presence of a more or less continuous pat- 
tern of geographic variation in the specimens we 
assign to the Paraploactis kagoshimensis com- 
plex. The apparent continuity is not present in 
factors III—-VI and may be the result of distortion 
of the 2-space defined by the first two compo- 
nents, which explain but 36% of the total vari- 
ance. The overall amount of variation in the P. 
kagoshimensis complex is greater than that seen 
in other Paraploactis species and suggests the 
complex is composed of a heterogenous assem- 
blage of species (see subsection ‘‘Paraploactis 
kagoshimensis complex’’). 


KEY TO THE SPECIES OF PARAPLOACTIS 


la. Ventral surface of lower jaw smooth, not 
covered with prickles or papillose villi 
and with cirri almost entirely confined 
to outer margin (Figs. 3A, B) 
1b. Ventral surface of lower jaw covered with 
numerous prickles or papillose villi and 
with cirri on both inner and outer mar- 
gins (Figs. 3C, D) 
2a. Minute cirri on snout, those in larger 
patch on tip of snout and across lach- 
rymal bones about equal in size to 
those in smaller patch on nasal bones 
(Fig. 2). Fleshy pad on isthmus (Fig. 
3A) with its width about 4-4 of length 
P. trachyderma (Fig. 7) 
2b. Large, fleshy, and extensively branched 
cirri on snout; those in larger patches 


Ce 


411 


on tip of snout and across lachrymal 
bones very large, those in smaller patch 
on nasal bones smaller (Figs. 2, 8). 
Fleshy pad on isthmus (Fig. 3B) with 
its width about /2—%4 of length 
P. pulvinus (Fig. 8) 
3a. Fleshy pad present on isthmus (Fig. 3C), 

its width about 4-4 its length ...... 
P. intonsa (Fig. 9) 
3b. No fleshy pad on isthmus (Fig. 3D) 

‘*P. kagoshimensis complex’’ (see text) 


Paraploactis trachyderma Bleeker 
(Figures 1, 2, 3C, 4C, 5-7; Tables 1, 2, 4) 


Paraploactis trachyderma BLEEKER, 1865:169-170 (original 
description; type locality Australia). BLEEKER 1876a:300 (in 
description of genus). MACLEAY 1881:141 (mentioned as 
from Australia). MCCULLOCH 1915:272 (listed; senior syn- 
onym of Aploactis lichen). MCCULLOCH & WHITLEY 
1925: 164 (listed; Moreton Bay; senior synonym of Aploactis 
lichen). MCCULLOCH 1929:397 (listed). WHITLEY 1933:100— 
101, pl. xu, fig. 5 (listed; figured; from Queensland). WHI- 
TLEY 1954:29 (new record for South Australia, Cape Jervis). 
ScoTrT 1962:162—163 (in key; brief description; figure). WHI- 
TLEY 1964:57 (listed). MARSHALL 1965:435, pl. 48 (common 
name; brief description; Queensland, South Australia; fig- 
ured). SCOTT ET AL. 1974:184 (brief description; South Aus- 
tralia, Queensland; Western Australia record not this spe- 
cies). Scott 1976:205 (listed, Australia). 

Aploactis lichen DE Vis, 1884:461 (original description; type 
locality Moreton Bay, Queensland). 

Erosa australiensis BORODIN, 1932:90-91, pl. 2 (original de- 
scription; type locality Southport, Queensland; figured). 
WHITLEY 1933:101 (compared with P. trachyderma). 


MATERIAL.—RMNH 5892 (1, 114, holotype of P. trachy- 
derma), Australia, no other data. 

The remaining specimens from Queensland. QM I.11/75 (1, 
103, holotype of A. lichen), Dunwich, Moreton Bay, donated 
G. Watkins. MCZ 33032 (1, 119, paratype of Erosa austral- 
iensis), Southport. AMS 1.7733 (1, 107), Moreton Bay, ex- 
change, by J. D. Ogilby, 1906. AMS 1.12514 (1, 116), Moreton 
Bay, exchange Amateur Fisherman’s Association. AMS 
I.18673-001 (1, 112), Moreton Bay, presented J. Lewis, Dec. 
1973. AMS IA.6170 (1, 69.4), and AMS IA.6171 (1, 37.8), 
Lindeman I., presented M. Ward, 1934. AMS IA.6236 (1, 
106), Cole I., near Lindeman I., presented M. Ward, 1934. 
AMS IB.6346 (1, 86.2), Moreton Bay, presented R. J. Smith, 
1963. AMS IB.7567 (2, 47.3-102), Moreton Bay, 1966. ANSP 
98678 (1, 98.5), Moreton Bay, T. C. Marshall, 12 Feb. 1952. 
FMNH 44968 (1, 89.2), Deception Bay, 27 Oct. 1952. QM 
1.382 (1, 105), Moreton Bay, G. Mindham, 2 Apr. 1912. QM 
1.467 (1, 86.5), Miora Banks, Harris and J. D. Ogilby, 28 May 
1912. QM I.510 (1, 92.5), Moreton Bay, J. Bancroft, 4 June 
1912. QM 1.4733 (2, 25.0-97.4), Bowen, E. H. Rainford, 5 
June 1931. QM I.12349 (1, 103), Moreton Bay, T. Marshall, 
24 Oct. 1950. QM I.12350 (1, 108), Moreton Bay, 18 Apr. 1953. 
QM 1I.2674—5 (1, 111), Moreton Bay. USNM 176933 (1, 83.9), 
Great Barrier Reef, 8 Apr.—29 May 1952, Col. Howard. UW 
7289 (1, 96.8), Moreton Bay, T. Marshall, Oct. 1950. 


412 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


A intonsa 


kagoshimensis 
complex 


pulvinus 


@ trachyderma 


“QUEENSLAND 


M@ SULU SEA 


@ FUJI 


Qe 
e @ 
@ 
HM HONG KONG 
m@ SRI LANKA e 
8 


FIGURE 6. 


Results of a principal components analysis using 45 morphometric and meristic variables on specimens of the 


genus Paraploactis. Above: first and second major axes. Facing page: first and third major axes. (See text for discussion, 


variables; Tables 4~7 for data.) 


DISTINGUISHING FEATURES.—(Measure- 
ments and counts are summarized in Table 4, 
body shape and color pattern in Fig. 7.) Ventral 
surface of lower jaw nearly smooth, with few 
papillose villi or prickles; about 5S—11 fleshy cirri 
in row along outer margin; about 0-4 fleshy cirri 
in row along inner margin; fourth left cirrus (out- 
er row) with 1+ papillae. Fleshy pad on isthmus 
about 4-3 as wide as long (Fig. 3). Numerous 
minute stubby cirri on snout in two patches, 
those of larger patch across tip of snout and 
lachrymal bones about equal in size to those in 
smaller patch on nasal bones (Fig. 2). Vertebrae 
26-28, usually 27. Dorsal fin rays XIII-XIV,8— 
10, usually XIV,9. Anal fin rays I,7-9, usually 
I,8. Live coloration unknown. Color in preser- 
vative (Fig. 7) brown with indistinct blotches 
and spots of darker and lighter brown on body 


and fins. Black spot at base of last dorsal rays. 
Poorly defined, nearly white spot or blotch sub- 
terminally on upper rays of pectoral fin. About 
6-10 dark brown bars radiating from orbit, par- 
ticularly noticeable on ventral margin (Bleeker 
(1865) states these bars were rosy in color on his 
specimen). Several specimens with a nearly 
white snout. 

DISTRIBUTION.—The type locality of Para- 
ploactis trachyderma was given by Bleeker only 
as Australia. Subsequent records indicate this 
species is apparently confined to eastern Aus- 
tralia (Fig. 1). While most records have come 
from Moreton Bay, Queensland, this species is 
known to occur southward to Southport and 
northward at least to Bowen and the vicinity of 
Lindeman Island. Whitley (1954:29) reports a 
specimen (SAM F.2659) taken on a weedy bot- 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 413 


tom at 20 m near Cape Jervis, South Australia, 
but this specimen could not be found for confir- 
mation. Otherwise the most southerly occur- 
rence for this species is Southport, Queensland. 
Other than Whitley’s record (possibly not this 
species), no depth of capture information is 
available. An old collection note accompanying 
one specimen (AMS IA.6236) states, ‘‘This spe- 
cies is very sleepy and lies in cranies in dead 
coral boulders. Grunts when held in hand.”’ 


Paraploactis pulvinus Poss and Eschmeyer, new 
species 

(Figures 1-6, 8; Tables 1-3) 

No literature applies to this species. 


MATERIAL.—Holotype: WAM P25528 (105 mm), Western 
Australia, Exmouth Gulf, CHALLENGER, trawl, 23 May 1971. 
Paratypes: All from Western Australia. WAM P20123 (1, 


119), Dampier Archipelago, NE Rosemary I., SE Elphics 
Knob, R. J. McKay, J. Stewart, A. Page, 5 Nov. 1971. CAS 
39680, formerly WAM P24908 (1, 111), Exmouth Gulf, 
trawled, Aug. 1974. QM I.12410 (1, 103), Exmouth Gulf, 
‘‘Gutters Area,’’ R. Rowe, trawled, 18.3 m, mud-silty bottom. 
WAM P23735 (1, 93.0), Exmouth Gulf, R. Rowe, Aug. 1973. 


Dr1aGnosis.—Dorsal XIII-XIV, 9-10. Anal I, 
8-9 (one abnormal specimen with I,4). Ventral 
surface of lower jaw smooth, without villi, 6-11 
fleshy cirri confined to row along outer margin. 
Large fleshy pad on isthmus about /2—4 as wide 
as long (Fig. 3). Snout very steep (Fig. 8) with 
numerous elongate fleshy cirri (Fig. 2); those in 
larger patch across tip of snout and lachrymal 
bones longer than those in smaller patch on na- 
sal bones. Lachrymal bone (infraorbital 1) with 
posterior two spines widely divergent; anterior- 
most spine very small. 


414 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


FIGURE 7. 


DESCRIPTION.—(Counts and measurements 
summarized in Table 3, body shape and color 
pattern in Fig. 8; based on six specimens.) A 
species of Paraploactis with features given un- 
der the genus above. Pectoral rays 14. Vertebrae 
27. All gill rakers very short, stubby, total 7-11, 
0-3 on upper arch, 7-9 on lower arch. Lachry- 
mal bone (infraorbital 1) with three spines, an- 
teriormost as inconspicuous lump, posterior two 
with points widely divergent (Figs. 7-8). Heav- 
ily ossified knobs, spines, and ridges on head, 
particularly in interorbit. Lateral line scales 12- 
14. Ventral face of lower jaw smooth, fleshy, 
without villi; 6-11 fleshy cirri in row along outer 
margin. Cushionlike pad on isthmus large, about 
Y2-34 as wide as long, abutting fleshy posterior 
border of lower jaw anteriorly so as to form a 
disclike pad on ventral side of head (Fig. 3). 
Large tuft of elongate fleshy cirri across nasal 
bones, on lower cheek anterior to lowermost 
preopercular spine, and a few near proximal end 
of branchiostegals. Papillose cirri on pectoral fin 
and on spines of dorsal fin, very large on first 
three spines. 

Color in life unknown. Color pattern in pre- 
servative, though variable, about as in Figure 
8. Body buff to tan with darker brown mark- 
ings in irregular patches or bands. Irregular 
bands or patches of black on fins. Pectoral fins 
whitish distally. Skin appears to be sloughing off 


Lateral view of Paraploactis trachyderma (AMS 1.12514, 116 mm). 


in two specimens (WAM P23735 and CAS 
39680). 

DISTRIBUTION.—Paraploactis pulvinus has 
been taken by trawling from Exmouth Gulf and 
Rosemary Island in the Dampier Archipelago 
and at Hampton Harbour, Western Australia 
(Fig. 1). At the first locality, one specimen (QM 
1.12410) was trawled on a mud-silty bottom at 
a depth of 18.3 m. At Hampton Harbour a spec- 
imen (CSIRO 2429) was taken in a prawn trawl 
at a depth of 5.5 m. 

ETYMOLOGY.—From the Latin pulvinus, a 
pillow, in reference to the fleshy pad upon which 
this fish rests its head. 


Paraploactis intonsa Poss and Eschmeyer, new 
species 
(Figures 1-6, 9; Tables 1, 2, 5) 


Paraploactis trachyderma (not of Bleeker): MEES 1960:20 (as 
new record for Western Australia; brief description [WAM 
P4714)). 


MATERIAL.—Holotype: WAM P12100 (125 mm) Western 
Australia, Shark Bay, W. and W. Poole, Sept. 1963. 

Paratypes: All from Western Australia. WAM P25840—001 
(1, 128), Shark Bay, R. McKay. WAM P8923 (1, 126), Shark 
Bay, E. Barker, Oct. 1964. WAM P5874 (1, 121), Shark Bay, 
W. and W. Poole, July 1963. CAS 39599, formerly WAM 
P5378 (1, 118), Shark Bay, W. and W. Poole, June—July 1961. 
WAM P23605 (1, 114), Carnarvon, 24 m, dredged, P. Heald, 
19 July 1972. WAM P4714 (1, 112), Shark Bay, Western Bay, 
R. J. McKay, July 1958, WAM P6069 (1, 111), Shark Bay, R. 
J. McKay. CAS 39600, formerly WAM P5388 (1, 110, cleared 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 415 


FiGuRE 8. Lateral view of Paraploactis pulvinus (holotype, 105 mm). 


and alizarin stained) and WAM P14683 (1, 97.5), Shark Bay, 
R. J. McKay. 

DIAGNosIs.—Dorsal XIV—XV,9-11, usually 
XIV,10. Anal I,9-10, usually I,9. Ventral sur- 
face of lower jaw with numerous villi and cirri 
on inner and outer margins. Fleshy pad on isth- 
mus about 14-4 as wide as long (Fig. 3). Snout 
with numerous cirri (Figs. 3, 9). 

DESCRIPTION.—(Counts and measurements 
summarized in Table 6; body shape and color 
pattern in Figure 9; based on ten specimens.) A 
species of Paraploactis with the features given 
under the genus above. Pectoral rays 14. Ver- 
tebrae 27-28, usually 28. All gill rakers very 
short, stubby; total 6-9; 0-2 on upper arch, 6— 
8 on lower arch. Lachrymal bone (infraorbital 
1) with three spines, anteriormost small, poste- 
rior two larger, with points moderately diver- 
gent. Lateral line scales 12-15. Ventral surface 
of lower jaw with numerous villi and papillose 
cirri, 8-11 cirri in row along outer margin, 4-8 
cirri in row along inner margin; fourth left cirrus 
(outer row) with 1-11 papillae. Cushionlike pad 
on isthmus about “4-4 as wide as long (Fig. 3). 
Band of papillose cirri across tip of snout and 
lachrymal bones. Papillose cirri over nasal 
bones, lower preopercular spines, pectoral fin, 
and dorsal spines; very large cirri on first three 
dorsal spines. Gasbladder masculature inserting 


on parapophyses of vertebrae 6—11 [CAS 39399, 
formerly WAM P5378]. 

Body color in preservative white to tan with 
blotches or irregular bands of brown. Dark 
brown or black mottling on fins. The skin ap- 
pears to be sloughing off in several specimens. 

REMARKS.—Dr. R. J. McKay has kindly pro- 
vided us (in litt., 13 July 1977) with the following 
additional information from field notes on spe- 
cies of Paraploactis taken during a trawling sur- 
vey in Shark Bay in 1960. We assume these re- 
marks apply to this species. 

‘*Bottom was usually sand and silt, depths 7- 
9 fathoms. First dorsal spines capable of being 
locked erect and spines with posterior groove. 
When preorbital spine is locked erect this action 
withdraws the eye into the orbit. After freezing 
specimens the mucous layer may be rubbed off 
to reveal mottled coloration of the skin. Life col- 
ours were greyish, brownish; mottled with grey, 
pink and brown; pale grey blotched with pink or 
bright orange; purple-brown with pink or orange 
blotches; mostly orange with grey blotches; one 
specimen orange with some pink blotches below 
eye. Two mature females, total lengths 140 and 
160 mm, running ripe on September 18, 1960. 
Eggs are small and orange in colour. In May, 
1960 all females (14) were spent and contained 
white grey silt in gut.” 


416 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


FIGURE 9. 


DISTRIBUTION.—Paraploactis intonsa is 
known from Western Australia in the immediate 
vicinity of Shark Bay (Fig. 1), in 7-24 m. 

ETYMOLOGY.—From the Latin intonsa, 
bearded or unshaven, in reference to the nu- 
merous cirri on the lower jaw and snout which 
give the face a somewhat shaggy appearance. 


Paraploactis kagoshimensis Complex 


DISTINGUISHING FEATURES.—Dorsal fin rays 
XIII-XIV ,9-10; total 23-24. Anal fin rays I,8-9. 
Numerous minute stubby papillae on snout. 
Ventral surface of lower jaw with numerous pa- 
pillae; 8-12 cirri in row along outer margin of 
mandible; 4—S cirri in row along inner margin of 
mandible. Fourth cirrus in outer row with 3-12 
papillae. Isthmus without a fleshy pad, though 
ending anteriorly in a small fleshy lump (Fig. 
3D). 

REMARKS.—The Paraploactis kagoshimensis 
complex is composed of four nominal species: 
Tetraroge kagoshimensis Ishikawa, 1904, in 
Jordan and Starks; Cocotropus obbesi Weber, 
1913; Aniculerosa taprobanensis Whitley, 1933; 
and Cocotropus hongkongiensis Chan, 1966. 
Because of the rarity of specimens, the status of 
these nominal species is uncertain. Judging from 
the specimens available to us, the complex prob- 
ably represents at least four species, though a 
hypothesis of marked regional geographic dif- 
ferentiation within a single wide-ranging species 
can not as yet be rejected. As discussed above 


Lateral view of Paraploactis intonsa (holotype, 125 mm). 


(Principal Components Analysis), the amount of 
variation seen in this group is greater than that 
seen in other species of Paraploactis. That sev- 
eral species are represented is further substan- 
tiated by the failure of a single-linkage clustering 
attempt over all specimens to group represen- 
tatives of this complex as a single unit. The re- 
sults of another single-linkage clustering attempt 
in which a ‘“‘city-block metric’’ is also used as 
a measure of phenetic distance and from which 
specimens not belonging to the Paraploactis ka- 
goshimensis complex have been excluded are 
presented in Figure 10. We present below the 
features which we suspect corroborate a multi- 
species hypothesis for the complex, and where 
possible, attempt to assign the specimens avail- 
able to us to existing names. 


Paraploactis kagoshimensis (Ishikawa) 
(Figures 1, 6, 10, 11; Tables 1, 2, 6) 


Tetraroge dermacanthus (not of Bleeker): ISHIKAWA & MAT- 
SUURA 1897:50 (listed; Kagoshima, Japan). 

Tetraroge kagoshimensis ISHIKAWA in JORDAN & STARKS, 
1904:171 (original description; type locality Kagoshima, Ja- 
pan; see remarks below). ISHIKAWA 1904:13-14, fig. 1 on 
pl. 7 (description). 

Erisphex kagoshimensis: JORDAN & STARKS 1904:169, 171, 
175 (placed in new genus Erisphex with type-species Co- 
cotropus pottii Steindachner; in key; name and descmrption 
from Ishikawa 1904; see remarks below). Jordan, Tanaka, 
& Snyder 1913:251 (listed: Kagoshima, Japan; common 
name Huurai-okoze). WEBER & DE BEAUFORT 1915:275—- 
276 (compared with Cocotropus dezwaani). 

Erisphex potti: TANAKA 1931:36 (in part, kKagoshimensis in 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 417 


BMNH 
CAS AMS CSIRO AMS 1933. 
5931 E.2944 A3237 1A.3951 8.12.50 


| 


+ 


4 


1) 
w 


(K= 


DISTANCE 
to) 


MINKOWSKI 


ZMA 


BMNH BMNH BMNH 
110 FRSKU 1862 1862 CMFRI 1965. 
-243 S364 11.1.55 11.1.56 141/511 11.6.4 


1.0 | 
FiGuRE 10. 


synonymy). KAMOHARA 1952:69 (in part, Kagoshimensis in 
synonymy). KAMOHARA 1964:75 (in part, kKagoshimensis in 
synonymy). 

Cocotropus kagoshimensis: MATSUBARA 1943:467—-470, in 
keys and figs., on earlier pages, fig. 155 (osteology; in keys; 
description of a specimen from Kagoshima). MATSUBARA 
1955: 1097 (in key; brief description). SHINO 1972:178 (com- 
mon name Rough Velvetfish). 


NOMENCLATURAL REMARKS.—Jordan and 
Starks (1904:175) quoted Ishikawa’s manuscript 
description of this species, used the scientific 
name selected by him, and cited him as the au- 
thor of the species. They provided brief infor- 
mation about the species, included it in a key, 
and referred it to their new genus Erisphex. We 
believe that Ishikawa (in Jordan and Starks) was 
the person responsible for the ‘‘validating con- 
ditions’ (see Article 50c, International Commis- 
sion on Zoological Nomenclature, 1964) and was 
responsible, although perhaps not alone, for the 
‘‘name and the conditions that make it avail- 
able’’ (see Article 50a). This interpretation par- 
allels the suggestion made by Sabrosky (1972, 
1974) that the ‘‘conditions that make it [name] 
available’ are the conditions other than publi- 
cation. Ishikawa’s manuscript was, however, 


Phenogram of computed relationships among specimens of the P. kagoshimensis complex. (See text.) 


published 17 days after that of Jordan and 
Starks. The dates of publication seem reliable. 
The Jordan and Starks paper is listed in the table 
of contents of volume 27 of the Proceedings of 
the United States National Museum for 1904 
with January 22, 1904, stated as the date of pub- 
lication. On the inside back cover of Ishikawa 
(1904) the printing date is given as 5 February 
and the distribution date as 8 February 1904. We 
follow all previous authors, including Jordan and 
Starks, in recognizing Ishikawa as the author. 
The species should be cited as Paraploactis ka- 
goshimensis (Ishikawa, 1904, in Jordan and 
Starks). 


MATERIAL.—FRSKU $364 (1, 37.3), Japan, Kagoshima, M. 
Abe, no other data (specimen described by Matsubara 1943). 
BMNH 1862.11.1.55-56 (2, 71.1-85.1), Japan, purchase of 
Mr. Jamrach, no other data. 


DISTINGUISHING FEATURES AND REMARKS.— 
We include here the specimens listed above. The 
type of P. kagoshimensis apparently is lost. 
Counts and measurements are presented in Ta- 
ble 7 and body shape in Fig. 11. Unlike some of 
the other specimens referable to the P. kago- 
shimensis complex, these have the membranes 


18 


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POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 419 


FIGURE 14. 


of the spinous dorsal fin at the level of the sixth 
to tenth spines with the spine tips free of skin 
posteriorly. The dorsal spines are somewhat less 
flexible than in some other specimens. One Aus- 
tralian specimen (CSIRO A3237, see Paraploac- 
tis species) also shows these features. 


Paraploactis taprobanensis (Whitley) 

(Figures 1, 6, 10, 12; Tables 1, 2, 6) 

Aploactis aspersa (not of Richardson): JOHNSTONE 1904:202, 
219, pl. 1, fig. 5S (description; from south of Adam’s Bridge, 
Ceylon [Sri Lanka]; misspelled species name aspersa for 
Richardson’s aspera). 

Aniculerosa taprobanensis WHITLEY, 1933:101 (original de- 
scription; based on Johnstone 1904). MUNRO 1955:241, pl. 
48, fig. 5, (description; figure from Johnstone). 


MATERIAL.—CMFRI F.. 141/511 (1, 83.2), se. coast of India, 
Veldalai, Gulf of Mannar. 


DISTINGUISHING FEATURES AND REMARKS.— 
Whitley did not see the specimen which John- 
stone described. We have been unable to locate 
this specimen. Alwyne Wheeler (in litt., 31 Aug. 
1975) has kindly informed us that both John- 
stone and Herdman were associated with the 


Lateral view of Paraploactis hongkongiensis (holotype, 95.2 mm). 


University of Liverpool, and following Herd- 
man’s resignation, his entire collection was dis- 
posed of by his successor. It seems likely that 
the specimen was lost at that time. 

The specimen we list above fits the descrip- 
tion given by Johnstone and is from the same 
general locality. The specimen differs from all 
others of the complex in the notably shorter 
lengths of the dorsal fin spines, and in lacking 
vomerine teeth. In other respects it is similar to 
P. kagoshimensis. 


Paraploactis obbesi (Weber) 
(Figures 1, 6, 10, 13; Tables 1, 2, 6) 


Coccotropus obbesi WEBER, 1313:503-504, figs. 104-105 
(original description; type locality North Ubian I., Sulu Ar- 
chipelago [generic name misspelled]). HERRE 1952:462—-464 
(description from Weber; in key; generic name spelled Co- 
cotropus on p. 462). HERRE 1953:575 (listed). 

Membracidichthys obbesi: WHITLEY 1933:102 (as the type- 
species of new genus; brief generic description). 

Cocotropus abbesi: MATSUBARA 1943:472 (compared with 
Cocotropus masudai; specific name misspelled). 

Erisphex obbesi: WEBER & DE BEAUFORT 1962:59-63, fig. 13 
(description; in key; figure from Weber 1913). 


— 


FiGures 11-13. 


Fig. 11. Lateral view of Paraploactis kagoshimensis (BMNH 1862.11.56, 85.1 mm). Fig. 12. Lateral view 


of Paraploactis taprobanensis (CMFRI F.141/511, 83.2 mm). Fig. 13. Lateral view of Paraploactis obbesi (holotype, 38.5 mm). 


420 


MATERIAL.—ZMA 110.243 (38.5, holotype of Coccotropus 
obbesi), Sulu Archipelago, anchorage off North Ubian I., 
06°7.5'N, 120°26'E, 16-23 m, Lithothamnion bottom, M. We- 
ber, SIBOGA sta. 99, 28/30 June 1899. 


DISTINGUISHING FEATURES AND REMARKS.— 
The dorsal spines in this nominal species are 
somewhat more flexible than they are in other 
specimens of the complex and notably curved 
rearward at their tips, and the dorsal fin mem- 
branes at the level of the sixth to tenth spines 
are not free of skin posteriorly (Fig. 13). In these 
respects, it resembles a specimen of the complex 
taken from Fiji (CAS 5931) but from which it 
differs in number of pectoral rays (13 in the type, 
14 in the Fiji specimen), an almost constant 
character in other species of Paraploactis. 


Paraploactis hongkongiensis (Chan) 
(Figures 1, 6, 10, 14; Tables 1, 2, 6) 
Cocotropus hongkongiensis CHAN, 1966:12—16 (original de- 


scription; type locality, Sharp I., inside of Port Shelter, 
Hong Kong). 


MATERIAL.—BMNH 1965.11.6.4 (95.2, holotype of Coco- 
tropus hongkongiensis), Hong Kong, West Shore of Port Shel- 
ter, the New Territories, 0.6-2.1 m, rotenone, bottom of 
rocks, boulders, and scattered patches of algae and sand, J. 
D. Bramhall. 


DISTINGUISHING FEATURES AND REMARKS.— 
This presumed species is known only from the 
holotype (Fig. 14). Measurements and counts 
are presented in Table 7. It is the largest speci- 
men referable to this complex. The dorsal spines 
are flexible, with the spine tips curved rearward. 
The dorsal spine tips are not free of membrane 
posteriorly. The membranes between dorsal 
spines at the level of spines 5-10 are deeply in- 
cised, more so than in P. obbesi. Also in com- 
parison with P. obbesi, P. hongkongiensis ap- 
pears to have a deeper body, especially 
posteriorly, more dark areas on the body, and 
a more compressed head with a long snout and 
more upturned mouth. 


Paraploactis species 
(Figures 1, 2, 6, 10; Tables 1, 2, 6) 


Aploactis milesii (not of Richardson): SEALE 1935:365 (brief 
description; from Suva, Fiji Islands, [CAS 59317]). FOWLER 
1959:357-358 (in part; one record from Fiji based on Seale 
1935). 


MATERIAL.—AMS IA.3951 (1, 57.7) Queensland, off Gat- 
comb Head, Port Curtis, 16.5—18.3 m, M. Ward & W. Board- 
man, July 1929. AMS E.2944 (1, 82.6), Queensland, 22.5 km 
SE of Cape Capricorn, 23.8 m, ENDEAvouR, 29 July 1910. 
BNMH_ 1933.8.12.50 (1, 39.0), Queensland, Great Barrier 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


Reef, 0.4 km N of North Direction I., Great Barrier Reef 
Expedition. CSIRO A327 (1, 15.7), Queensland, Gulf of Car- 
pentaria, RAMA sta. 158, 16°46.4’S, 140°25.0’E, 18.3 m, trawl, 
1S Sep. 1963. CAS 5931 (1, 42.1), Fiji Islands, Vitti Levu, 
Suva, 24 Apr. 1933. 


DISTINGUISHING FEATURES AND REMARKS.— 
The material listed above constitutes a rather 
heterogeneous group encompassing nearly as 
much variation as seen among the previously 
considered nominal species and probably rep- 
resents more than one species. Some specimens 
appear similar to those from Japan, while others 
are similar to the specimen from the Sulu Sea. 
We are unable to assign them with certainty to 
any of the nominal species discussed above. 


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Bull. Zool. Nomencl. 29(2):79-91. 

1974. Article 50 and questions of authorship. 
Z.N.(S.)1925. Bull. Zool. Nomencl. 31(4):206-208. 

Scott, E. O. G. 1942. Observations on some Tasmanian 
fishes. Part V. Paps. Proc. Roy. Soc. Tasmania 1941:45—54, 
jolle 7 

. 1976. Observations on some Tasmanian fishes: Part 
XXII. Paps. Proc. Roy. Soc. Tasmania 110:157—217. 

Scott, T. D. 1962. The marine and fresh water. fishes of 
South Australia. W. L. Hawes, Government Printer, Ade- 
laide. 338 pp., figs. 


422 


,C. J. M. GLOVER, AND R. V. SouTHcoTT. 1974. The 
marine and freshwater fishes of South Australia. (Second 
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392 pp., figs. 

SEALE, A. 1935. The Templeton Crocker Expedition to west- 
ern Polynesian and Melanesian islands, 1933, no. 26. Fish- 
es. Proc. Calif. Acad. Sci., Ser. 4, 26(27):337-378, pls. 20- 
23% 

SHINO, S. M. 1972. List of English names of Japanese fishes 
with proposition of new names. Sci. Rep. Shima Marineland 
1:1-210, 1 pl. 

SmiTH, J. L. B. 1935. New and little known fishes from South 
Africa. Rec. Albany Mus. Grahamstown 4:169-235. 

. 1958. Fishes of the families Tetrarogidae, Caracan- 
thidae and Synanciidae from the Western Indian Ocean, 
with further notes on scorpaenoid fishes. Ichthyol. Bull. 12: 
167-181, 2 pls. 

STEINDACHNER, F. 1896. Bericht tiber die wihrend der Reise 
Sr. Maj. Schiff “‘Aurora’’ von Dr. C. Ritter V. Microszews- 
ki in den Jahren 1895 und 1896 gesammelten Fische I. 
Fische von Kobe, Hiogo und Nagasaki. Ann. Naturhist. 
Hofmusems Wien 11(12):197—230, 4 pls. 

TANAKA, S. 1915. Ten new species of fish from Japan. [In 
Japanese] Dobutsugaku Zasshi [Zool. Mag.], Tokyo 
27(245):565—568. 

. 1918. Figures and descriptions of the fishes of Japan 

including Riukiu Islands, Bonin Islands, Formosa, Kurile 

Islands, Korea, and southern Sakhalin. Mauren Co., Ltd., 

Tokyo 18:495-514, pls. 136-137. 

1931. On the distribution of fishes in Japanese 
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TEMMINCK, C. J., AND H. SCHLEGEL. 1843. Pisces. Jn Fauna 
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Siebold. Conjunctis studiis C. J. Temminck et H. Schlegel 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


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THOMPSON, J. M. 1967. New species and new records of fish 
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Waite, E. R. 1903. Additions to the fish fauna of Lord Howe 
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. 1921. Illustrated catalogue of the fishes of South Aus- 

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. 1952b. Some noteworthy fishes from eastern Austra- 

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13:235-243, 2 pls., 5 figs. 


32. 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 


TABLE 3. COUNTS AND MEASUREMENTS OF SPECIMENS OF Paraploactis pulvinus. 


Holotype 
WAM WAM CSIRO WAM QM WAM 
P25528-001 P29123 C2429 P24908 1.12410 P23735 
Standard length 105 119 95.9 111 100 93.0 
Dorsal rays XIV,9 XIV,10 XIV.9 XIV,9 XIII,10 XIII,11 
Anal rays 1,8 I) 1.4 1,8 9 189 
Pectoral rays 14,14 14,14 14,14 14,14 14,14 14,14 
Gill rakers (left) 14+8=9 1+7=8 3+7=10 0+7=7 2+6=8 1+6=7 
Gill rakers (right) 2+9=11 1+8=9 3+8=11 0+8=8 1+6=7 2+6=8 
Lateral line scales P2113) 14,14 14,14 14,12 12,13 13,14 
. Vertebrae 244] 27 iT, Di) 27 Di 
Head length 41.0 44.9 35.7 43.6 40.3 37.8 
Snout length 18.9 NA) 13.4 16.2 14.7 14.6 
. Orbit diameter 7.0 8.4 7.6 8.3 7.4 ell 
. Interorbital width 13.5 14.9 12.0 igh 12.4 Hite 
. Jaw length 14.0 1522 13 15.3 14.2 13.0 
. Postorbital length 19.1 22.2 16.5 Die 19.5 18.1 
. Body depth 39.1 43.4 35.0 44.2 40.6 36.9 
. Predorsal length 27.9 29.3 24.6 PLD) 26.6 23.6 
. Anal fin length 3559) 41.7 23.6 39.4 B55. 30.8 
. Caudal fin length 7773.3} 30.4 19.3 27.8 21.8 222 
. Pectoral fin length 21.8 26.5 23.2 29.0 Darl 26.6 
. Pelvic fin length 18.8 PY 19.0 20.3 17.0 18.0 
. First dorsal spine length 13.2 13.1 10.4 1Be2 14.3 12.6 
. Second dorsal spine length 13.8 15.9 1222 14.2 14.2 14.0 
. Third dorsal spine length 12.0 14.7 12a 14.2 13357/ 12.6 
. Fourth dorsal spine length 12.0 12.0 8.0 10.6 9.8 Onl 
. Fifth dorsal spine length 12.0 95 8.1 12.2 10.3 9.4 
. Penultimate dorsal spine length 12.0 12.9 11.6 iil? 10.9 9.3 
. Last dorsal spine length 11.3 13.0 11.8 11.0 Wil OFF, 
. First anal spine length 4.9 6.9 6.6 8.7 SY 4.3 
. Least depth of caudal peduncle 13.6 16.3 eS IS} 7/ 13.3 11.5 
. Snout to base of 2nd dorsal spine 30.4 32.8 26.8 31.0 30.1 28.2 
. Snout to base of 3rd dorsal spine 31.8 34.6 28.4 Bits 31.3 30.3 
Snout to base of 4th dorsal spine 39.8 42.5 36.0 42.0 41.5 38.3 
. Snout to base of Sth dorsal spine 44.7 49.7 40.0 48.4 46.4 43.9 
. Distance between interorbital ridges 7.0 8.5 7.8 3 Ue 7.4 
. Papillae per cirrus 0 0 0 0 3 | 
. Cirri along outer margin of lower jaw 11 11 6 10 12 8 
. Cirri along inner margin of lower jaw 0 0 0 0 0 0 
. Pad length 10.5 9.6 9.0 11.8 12.7 WS) 
. Pad width 7.9 8.9 Weil 9.6 10.0 7.0 
. Length of cirri on snout 3.3 4.8 1.8 3.4 4.9 2.4 
. Length of cirri on nasals 4.8 3h5) 1.3 5.3 2.0 1.9 
. Interorbital pit depth 32 2.8 2.8 3.2 Bw 3.0 
. Width Ist dorsal spine at midlength 2.7 3.8 3.0 3.5) 38) 7451 
. Incision of dorsal fin membrane at 
fourth spine (from tip to membrane) Tiel ies 7.0 6.3 S)// 7.8 


424 


TABLE 4. 
listed in Table 3.) 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


Holotype 
RMNH QM 
5892 111/75 
Ir 94.8 103.4 
2 XIII,10 XIV,9 
3. 1,8 iS) 
4. 14,14 14,14 
3. 1+6=7 1+7=8 
6. 1+7=8 1+7=8 
ik 15,14 13,12 
8. 27 27 
2). 94.8 41.6 
10. 15.4 14.3 
Il. 6.7 6.0 
2s 10.4 10.5 
13. 13.2 13.3 
14. 18.3 19.0 
ilSy BIS 35.0 
16. 24.5 24.2 
17. PS 29.1 
18. 18.5 D2 
19. 17.5 22.8 
20. 14.5 15.8 
PAE 10.4 G7 
22: 10.9 12.6 
3K. 3 11.0 
24. 7.0 7.0 
25 73 ed 
26. Tell 7.6 
27. 8.2 8.3 
28. 3),7/ 4.4 
29. 10.9 11.0 
30. 27.0 27.8 
Si: Maffei?) 30.5 
325 38.4 3955) 
33% 44.0 45.7 
34. 7.0 Wes) 
35: 1 2 
36 6 10 
37 0 3 
38. 5.4 ed 
39) 3}3) 3.9 
40. = 1.2 
41 1.0 0.4 
42 2.4 Zell 
43 Iles) 3.0 
44 4.7 5.6 


MCZ AMS QM SU 
33032. ~=—«1A.6236~—Ss«14733 20393 
102.8 106.5 97.4 90.0 
XIV.9 XIV.8 XIV,9 XIV,9 
1.8 1,7 1.8 1.8 
13,14 14,14 13,13 14,14 
1+8=9 14+7=8 14+6=7 0+6=6 
3+5=8  14+6=7 246=8  0+7=7 
12,13 13,13 14,14 15,15 
28 27 27 27 
39.3 42.9 40.0 34.1 
15.6 16.8 16.4 11.6 
7.0 7.5 5.9 6.1 
10.8 10.8 10.8 9.6 
13.1 14.6 13.4 11.8 
19.4 20.3 19.5 17.2 
37.1 35.1 32.3 31.4 
24.8 25.3 21.9 21.2 
31.8 33.6 32.3 28.8 
22.8 24.6 1238 19.9 
22.9 20.3 21.4 18.1 
18.5 18.1 15.6 16.7 
11.9 10.9 iia 10.0 
12.4 13.5 12.7 11.4 
12.3 hd7 10.8 10.7 
5.2 7.0 72 6.8 
5.6 7.6 7.4 7.0 
9.0 9.0 8.9 7.1 
9.1 9.1 10.5 7.4 
238 5.8 5.0 5.0 
11.7 11.9 12.1 10.7 
26.6 28.9 26.9 24.0 
29.1 31.3 30.3 26.3 
39.5 41.9 39.2 32.8 
43.2 46.1 44.6 37.8 
5.6 6.6 7.6 5.8 
I I 2 2 
8 5 10 10 
0 4 2 2 
6.4 7.4 es 5.5 
3.4 3.4 3.4 2.2 
0.6 0.8 1.0 0.5 
0.8 1.3 0.6 0.8 
2.1 2.4 1.8 1.7 
1.6 4.0 3.3 as) 
6.2 6.3 al 4.9 


FMNH 


44968 
89.2 


XIV.10 


ie) 
14,14 
1+5=6 
1+6=7 
13,14 
27 
37.5 
14.1 
3)o3) 
9.1 
13.5 
N7/o7 
34.5 
2eS 
31.8 
19.6 
ZED 
14.0 
(il 72 
11.6 
10.8 
6.4 
7.4 
8.9 
8.3 
4.6 
11.6 
24.0 
26.7 


COUNTS AND MEASUREMENTS OF SPECIMENS OF Paraploactis trachyderma. (Numbers at left refer to characters 


AMS 


1A.6170 


69.4 


XIV,9 


1,8 
14,14 


1+9=10 
1+7=8 


13,13 
27 
26.4 
9.9 
5.0 
6.9 
10.3 
11.8 
22.8 
16.4 
23.6 
1523 
14.7 
12.2 
7.8 


AMS 
1A.6171 


37.8 
XIV,9 
1.8 
14.14 
1+7=8 
0+7=7 
14.14 
27 


Ww ~I 00 CON OW ~) 


WwWNN Ww 
mMn— CO OOMNMWe NIDA UWOAN~IOAIWY 


mAMOnnNn = nN 
oo 


Ow WwW 


QM 
1.4733 


25.0 
XIII,9 
1,8 
14,14 
2+5=7 
1+7=8 
14,14 


TABLE 5. 


Holotype Paratype 
WAM WAM 
P12100 =P25840-001 
125 128 
XIV,10 XVv,9 

1,10 I) 
14,14 14,14 
1+7=8 1+6=7 
1+6=7 0+7=7 
A512 13,14 
28 28 
47.5 49.3 
18.3 18.9 
7.8 7.9 
10.8 i13}.33 
16.2 17.0 
24.0 23.8 
45.8 46.1 
2-3} 31.9 
46.9 43.3 
29.1 28.8 
29.2 28.5 
21:9 24.3 
15.7 11.6 
16.0 11.8 
13.9 10.7 
10.2 8.6 
10.7 9.4 
33 10.3 
14.1 10.4 
7.1 5.4 
16.7 16.4 
32.0 36.3 
34.5 38.3 
47.3 45.3 
51.0 51.8 
7.0 Tell 
10 3 
12 12 
4 4 
Oy 9.0 
2.8 3.0 
1.6 0.5 
0.3 0.4 
2.4 2.5 
Ips) 23 
10.2 4.8 


POSS & ESCHMEYER: AUSTRALIAN VELVETFISHES 


Paratype 
WAM 
P8923 


126 
XIV,10 
1,9 
14,14 
0+6=6 
1+7=8 
14,15 
28 
47.3 
17.6 
7.0 
12.6 
16.0 
22.1 
42.8 
28.5 
46.9 
29.0 
29.3 
21.6 
14.4 
14.7 
1329 
9.2 
10.8 
11.1 
NES 
5.6 
14.6 
33.9 


Paratype 
WAM 
P5874 


121 
XIV,10 
1,10 
14,14 
1+7=8 
1+7=8 
12,12 
28 
47.2 
20.0 
8.0 
12.4 
16.1 
22.0 
42.3 
31.6 
46.3 
22.3 
20.9 
20.3 
11.8 
12.9 
11720 
8.7 
9.4 
11.3 
11.0 
6.3 
14.9 
S)5)5// 


425 


COUNTS AND MEASUREMENTS OF TYPE-SPECIMENS OF Paraploactis intonsa. (Numbers at left refer to characters 
listed in Table 3.) 


Paratype Paratype Paratype Paratype Paratype Paratype 
WAM WAM WAM WAM WAM WAM 
P5378 P23605 4714 P6069 P5388 P14683 
118 114 112 111 110 97.5 

XIV,11 XIV,9 XIV,10 XIV,10 XIV 10 XIV,10 

1,9 HIS) 1,10 1,9 IES) IES 
14,14 14,14 14,14 14,14 14,14 14,14 
2+6=8 2+7=9 1+6=7 1+6=7 0+7=7 0+7=7 
2+7=9 1+7=8 1+6=7 1+7=8 1+8=9 0+8=8 
113}, 115) 14,13 14,14 14,14 13,15 15,14 
28 28 28 27 28 27 
45.8 43.2 42.7 43.9 41.9 30.0 
17.8 17.4 17.5 7a 16.8 14.1 
8.0 7.0 6.7 7.6 7.6 7.6 
i) 11.9 7258) 12.8 10.8 10.2 
7 15.8 16.1 16.8 15.8 14.5 
Mai 72 20.1 20.7 Ae 20.9 18.3 
41.4 38.8 39.9 43.1 SES 36.3 
28.7 28.0 27.1 7), 25.9 24.4 
45.3 41.7 43.4 40.1 39.2 36.9 
28.7 26.5 27.4 24.6 23.6 23.0 
30.7 26.6 23.5 24.4 PDS) 26.5 
19.8 19.6 19.5 20.5 20.8 17.8 
14.8 13.3 11.4 ES 13.5 10.4 
14.0 13.4 11.8 12.0 14.1 10.7 
13.4 2 11.6 11.4 13.0 10.9 
all 10.5 9:9 8.0 7.0 7.0 
8.1 Wes) 8.9 8.1 6.5 Toll 
IWS7/ 11.2 10.7 11.1 10.3 Oo 
11.6 11.3 10.5 IES 10.0 9:9 
S57 6.7 6.5 5),8) 5.0 5:3 
15.7 12.6 14.9 113353 12.9 12.3 
32.0 Sie IVS) 32.0 29.6 27.9 
33.6 S357 31.6 33.3 31.6 29.4 
43.3 43.4 42.1 41.4 39.0 37.7 
49.0 49.4 47.5 47.5 44.0 41.7 
7.0 6.4 Wes} 8.3 6.7 6.1 
i 10 11 | 3 6 
10 13 11 12 8 12 
8 6 4 4 4 6 
6.8 8.0 6.9 7.6 8.9 6.8 
3.0 3.4 ell DS 3.6 2.5 
1.1 0.9 0.4 0.5 a2 0.2 
1.0 0.9 0.1 0.5 0.6 0.6 
2.4 1.6 DD, 1.6 Dal ja) 
7.8) Dal Dee 7,1 Wea 1.8 
6.5 8.7 5.8 5.5 4.0 5.4 


426 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 18 


TABLE 6. COUNTS AND MEASUREMENTS OF SPECIMENS OF THE Paraploactis kagoshimensis COMPLEX. (Numbers at left 
refer to characters listed in Table 3.) 


BMNH BMNH BMNH BMNH 
1862 1862 FRSKU CMFRI ZMA 1965 = AMS AMS 1933. CSIRO CAS 
1.1.55 11.1.56 $364. F.141/S11 110.243 11.6.4 —- TA.3951- E2944 8.12.50 A3237. ‘5931 
85.1 71.1 373 83.2 39.8 95.2 ST 82.6 39.0 80.0 42.1 
2 XHI,10 © XIN,10-XIV,10 © XT,10.--XTNI,10—-XTN,10 XIV,10-XI,10 = XV.9—XIV.10—XIV.9 
3 1,9 1,8 1,8 1.8 1.8 1,8 1,9 1,9 1,9 1,9 1,8 
4 13}135) 2133 a 135 13,13" 913313 MBI aa) 44) 71414 1 
5. 248=10 2+7=9 14+6=7 24+6=8 14+4=5 14+6=7 14+5=6 O+7=7 0+6=6 1+5=6 1+46=7 
6. 24+7=9 24+6=8 046-6 146=7 24+4=6 24+5=7 14+6=7 > 14+6=7 14+6=7 14+5=6 14+6=7 
7 14,13 15,14 42- 16,14" 42513 AA gat EP Math12e 9 1614 ne aoe 
8 27 27 28 27 27 27 28 27 27 28 27 
9. 30.5 24.8 13.3 28.8 13.9 34.9 20.8 32.1 14.6 28.4 15.0 
10. 10.2 7.5 4.2 10.3 4.8 13.2 1 11.9 4.1 9.9 5.4 
I. 6.4 5.4 au 5.5 2.8 7.0 5.1 6.2 3.5 5.4 3.6 
12. 6.9 5.8 3.2 6.9 3.9 8.6 5.8 7.8 4.1 7.0 4.2 
13. 10.8 8.6 5.0 10.6 4.8 12.8 el 12.1 Sai 11.2 5.9 
14. 14.0 11.5 5.7 13.5 6.3 15.6 9.2 14.0 6.6 13.2 6.7 
15. 25.9 19.4 11.7 25.9 12.0 33.4 17.9 28.0 12.1 26.3 13.3 
16. 17.9 9.2 7.3 16.0 7.6 21.6 12.5 19.3 Ta 16.9 9.1 
17. 25.3 23.6 12.4 24.8 13.5 33.4 20.4 27.0 13.0 25.3 14.5 
18. 18.1 16.9 9.1 16.3 9.0 28.8 14.0 20.1 9.0 18.1 11.4 
19. 17.5 16.2 9.1 19.9 7.7 23.8 12.9 22.0 9.8 18.7 8.3 
20. 13.2 12.4 i) 12.6 7.5 177 11.0 15.7 7.8 13.8 7.4 
i 11.0 9.3 4.6 8.8 4.6 9.7 9.1 13.6 6.3 10.7 5.1 
22: 12.8 9.4 5.3 9.8 5.5 12.7 10.0 14.6 7.2 11.5 5.3 
23. 11.0 9.4 4.9 9.2 52 7 9.7 14.6 6.2 11.7 4.9 
24. 6.6 6.8 3.8 4.5 Ds 9.2 4.9 8.8 3.6 1D 3.0 
25; 6.5 5.3 a2 4.5 2.8 6.9 5.3 8.3 3.3 5.6 3.1 
26. 7.8 7.0 3.3 6.6 4.3 9.2 6.9 8.2 3.9 9.8 4.1 
Dil 7.9 7.4 3.3 6.5 4.4 9.0 TS 8.3 3.9 10.2 4.6 
28. 4.7 3M 0.6 3.5 2.2 So 3.0 4.4 DS 3.3 2.7 
29 9.7 7.4 4.2 8.3 4.8 12) 6.8 10.2 4.3 8.9 5. 
30. 19.7 17.1 8.5 17.3 8.3 24.4 57 23.8 10.4 21.3 10.9 
31. 22.6 18.7 9.0 19.1 10.0 26.4 19.5 28.5 12.0 21.6 12.1 
32 28.9 2212 10.3 24.2 13.0 32.3 20.5 29.3 14.3 25.0 14.4 
33 33.2 29.8 11.5 31.3 14.1 36.7 22.7 35.4 16.6 30.5 16.8 
34 4.5 3.6 2,2 4.2 2.5 SW) 5.2 5.2 Bp Tee 2.1 
35, 4 7 7 8 8 0 6 6 9 3 7 
36. 8 9 8 12 12 8 12 10 10 10 8 
ai. 7 6 6 4 4 4 4 6 4 4 6 
38. 0 0 0 0 0 0 0 0 0 0 0 
39 0 0 0 0 0 0 0 0 0 0 0 
40 0.1 0.1 0.1 - 0.1 0.1 0.2 0.1 + 0.2 0.2 
41 0.1 0.1 0.2 0.1 = 0.1 0.1 0.3 - 
42 1.9 1.4 0.5 3 0.4 1.9 1.7 1.7 0.6 2.0 0.8 
43 1.2 0.9 0.3 1d 0.5 1.5 | 2.4 0.7 1.4 0.4 
44 2.2 Ash 1.5 2.3 1.3 4.6 1.8 3.6 1.2 2.1 0.9 


—— 


PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 
FOURTH SERIES 


Vol. XLI, No. 19, pp. 427-452; 10 figs. 


January 24, 1979 


A REVISION OF MYCTEROMYIINI (*“SGENUS MYCTEROMYIA”’ OF 
AUTHORS), A NEW TRIBE OF NEOTROPICAL 
HORSE FLIES (DIPTERA, TABANIDAE) 


By 


Sixto Coscaron 
Facultad de Ciencias Naturales y Mueso, La Plata, Argentina 


and 
Cornelius B. Philip 


Research Associate, Department of Entomology, California Academy of Sciences, 
Golden Gate Park, San Francisco, California 94118 


ABSTRACT: 


Genitalic analyses have confirmed that the group of primitive flies now assigned to the genus 


Mycteromyia Philippi, 1865, in southern South America is composite. A review of species heretofore in- 
cluded here reveals that these flies are to be redistributed taxonomically as follows: of 13 species recently 
included in the genus under the primitive tribe Scionini (subfamily Pangoniinae), 2 are excluded as discussed 
below and only 3 are questionably retained in the tribe, but in the revived (previously synonymized) genus 
Caenopangonia Kroeber, namely hirtipalpis (Bigot) (type-species), brevirostris (Philippi), and asper (Philip). 
The remaining 8 accepted species, plus 7 previously undescribed, are distributed in the new tribe Mycter- 
omyiini, also of subfamily Pangoniinae, among 3 genera: Promycteromyia new genus for P. philippii (Philip) 
(type-species, new combination), P. galbina new species (d, 2, Chile), P. derocerca new species (d, &, 
Chile), P. eriodes (Philip) (new combination), P. cinerascens (Bigot) (new combination), P. pechumani new 
species (6, 2, Chile), P. penai new species (6, 2, Chile), P. murina (Philippi) (new combination), and P. 
xantha new species (d, Chile); Mycteromyia for M. conica (Bigot) (type-species) and M. etcheverryae new 
species (¢, 2, Chile); and revived Silvestriellus for S. patagonicus Bréthes (type-species), S. schlingeri new 
species (d, 2, Argentina), S. martinezi (Barretto and Duret) (new combination), and S. flaviventris (Barretto 
and Duret) (new combination). Mycteromyia bejaranoi Barretto and Duret is newly synonymized with C. 
hirtipalpis (Bigot). Pangonia obscuripennis Philippi, assigned by Kroeber (1934) to Mycteromyia and retained 
there by Fairchild (1971), is not included here, nor is Mycteromyia robusta Kroeber; we considered neither 
to belong in this revised group. 


INTRODUCTION 


It has become evident that there is more su- 
pra-specific diversity than previously supposed 
in the genus Mycteromyia Philippi, a group of 
primitive flies occurring in southern South 
America. We now consider certain characters 
that were until recently considered common to 
the group to be of polyphyletic origin, such as: 
wide fronts in both sexes, elongate, basally 
hinged proboscides not armored for ‘‘biting,”’ 
long wings with isolated clouds and closed first 
posterior (R;) cells, and often produced, snout- 


like faces (from which Philippi derived his ge- 
neric name—‘‘nose-fly’’). 

Because of our discovered heterogeneity in 
the Mycteromyia group, we realized that any 
proposed systematic revision would be depen- 
dent on the identity of the nominate type-species 
of this genus. This presents a nomenclatural 
problem that only can be resolved by ultimate 
action of the International Commission on Zo- 
ological Nomenclature (ICZN). 

THE TYPE-SPECIES OF THE GENUS Myctero- 
myia.—Pangonia conica Bigot from Chile was 


[427] 


428 


the only species included in the original generic 
characterization by Philippi (1865), though he 
described several other species in the genus. 
Without further characterization except in a ge- 
neric key, Enderlein (1922) designated P. conica 
as the type-species of Mycteromyia. The same 
designation was repeated in the neotropical cat- 
alog of Kroeber (1934) and by Hack (1953) in 
discussing the genus in Argentina. 

This concept was inadvertently changed by 
Fairchild (1971:12) when he stated, ‘‘Type spe- 
cies, Pangonia conica Bigot [(JEnderlein, 
1922:340) = philippii Philip.’’ Philip (1958) de- 
scribed M. philippii (syn. M. conica sensu Phi- 
lippi, not Bigot) after discovering and confirming 
(1968) that Philippi had misidentified P. conica 
Bigot. As shown below, the real M. conica (Big- 
ot) was inexplicably redescribed as M. fusca by 
Philippi. Following ICZN Articles 67 and 70a, 
which cover misidentifications of type-species, 
the International Commission has been peti- 
tioned (Philip, 1977) to use its plenary powers 
to declare P. conica Bigot as the type-species of 
Mycteromyia Philippi, (not sensu Philippi = 
philippii Philip). Genitalic study has shown that 
the two concepts are not congeneric so that the 
potential consequences of the ICZN decision are 
not trivial. Pending Commission action on the 
above petition, we shall continue to use the 
type-species concept in use prior to 1971 as per- 
mitted by the Rules. 

Within this group of flies, we find that genitalic 
characters support retention of only three spe- 
cies in the heterogeneous tribe Scionini, i.e., 
hirtipalpis Bigot, brevirostris Philippi, and asper 
Philip, with their more conventional, nonpro- 
duced faces, and in males, nonbulbous termi- 
nalia. These have been reassigned to the revali- 
dated genus Caenopangonia Kroeber. Among 
the remaining known species of ‘‘Myctero- 
myia,’ plus others described below, we have 
found very peculiar genitalic characters. Some 
of them are more primitive than in most more 
highly evolved Tabanidae, and others approach 
those in the family Pelecorhynchidae, which 
merit their taxonomic separation at least at the 
tribal level. On morphological evidence, we be- 
lieve that this generalized group may resemble 
hypothetical ancestors of known tabanids. This 
group has some plesiomorphic characters, some 
of which also occur in certain other dipterous 
families, such as the pelecorhyncids. We con- 
sider the latter to have more primitive genitalic 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


characters than the more specialized, higher Ta- 
banidae. This opinion is reinforced by the re- 
stricted, relict-type of species distribution in 
southern Chile and Argentina. There are also 
apomorphic characters which facilitate separa- 
tion of different taxa. 

We are dividing the new tribe Mycteromyiini 
into the most generalized genus Promyctero- 
myia new genus, Mycteromyia Philippi, the 
type-genus, and the revived genus Silvestriellus 
Brethes. The order of textual treatment of taxa 
attempts to follow our concept of evolvement of 
characters from generalized to the more spe- 
cialized in different groups. 

We frequently use textual abbreviations for 
structures like T. IX, T. X, or St. VIII in ref- 
erence to respective tergites and sternites. ‘‘Soft 
abdomens”’ are those having less rigid scleroti- 
zation than usual. Symbols used in the figures 
are explained with initial usage (Figs. 1 and 2). 

Characters and symbols that relate to male 
genitalia are: in external (‘‘visible’’) view, the 
bulbous epandrium shield (Figs. 1B and IC, ep), 
cerci (Figs. 1B and 1C, ce); internal view by 
dissection, gonocoxite (Fig. 1D, go), with basi- 
style (ba) and dististyle (di), aedeagus (ae), apo- 
deme of aedeagus (aa), flagellar aedeagus (fa), 
and, in some taxa, very peculiar apical styli (st) 
and inner tuberosities (tu). Female genitalia: af- 
ter dissection, genital fork (Fig. 3G) with pecu- 
liar caudal ducts (cd) and ‘“‘sclerotized portions” 
(sc), sternite VIII (St. VIII), cerci (Fig. 1F, ce), 
tergites [IX and X (T. IX, T. X); T. IX with un- 
usual lateral expansions termed by us, lateral 
‘‘flaps’’ (If). Genitalic structures are thus dis- 
cussed from two aspects: external, in situ, and 
internal after dissection; ‘‘in slide view’’ refers 
to mounted preparations of internal structures. 


ACKNOWLEDGMENTS 


The authors are indebted for study material to 
many sources, but especially to Dr. L. L. Pechu- 
man, Cornell University, Ithaca, New York, 
Luis E. Pena G., Escuela Agronomica, and Dra. 
Maria Etcheverry, Centro de Estudios Ento- 
mologicos, both University of Chile, Santiago. 
We also acknowledge the U.S. Council for In- 
ternational Exchange of Scholars which spon- 
sored travel of the senior author to do this work, 
a substantial part of which was accomplished at 
the California Academy of Sciences, San Fran- 
cisco. The authors are also much indebted to 
Dr. P. H. Arnaud, Jr., and Mr. V. F. Lee, Cal- 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


FIGURE 1. 
appendages, lateral view. C. Epandrium and cerci in slide view. D. Gonapophyses and genital appendages. E. Spermathecal 
duct. F. Genital appendages. Abbreviations: aa, aedeagal apodeme; ae, aedeagus; ba, basistyle; ce, cercus; di, dististyle; Do, 
dorsal view, ea, endophallic apodeme; ep, epandrium; fa, flagellar aedeagus; go, gonocoxite; hi, hypoproct; If, lateral flap; pv, 
penis valve; sc, sclerotized portion; st, style; tix, tergite IX; tu, tuberosity; tx, tergite X; Ve, ventral view; vp, ventral plate of 
proctiger. 


ifornia Academy of Sciences, and Dr. R. S. 
Lane, California State Department of Health, 
Berkeley, for the considerable chore of review- 
ing the manuscript. 

Abbreviations of depositories of specimens 
are as follows: CAS—California Academy of 
Sciences, San Francisco; CEEUC—Centro de 
Estudios Entomologicos, Universidad de Chile, 
Santiago; LLP—Collection of Dr. L. L. Pechu- 
man, Ithaca; MACN—Museo Argentino de 


429 


Promycteromyia philippii (Philip). Male (A—D), female (E, F). A. Abdomen, dorsum. B. Epandrium and genital 


5 


Ciencias Naturales “‘Bernardino Rivadavia,’ 
Buenos Aires; MLP—Museo La Plata, La Plata; 
MNHN—Museo Nacional de Historia Natural, 
Santiago; MZSP—Museu de Zoologia da Uni- 
versidade de Sao Paulo; NMW—Naturhistor- 
isches Museum Wien. 


MYCTEROMYIINI, new tribe 


DIAGNosIs.—Species mostly grayish or yel- 
lowish to brown, thorax usually with four notal 


430 


stripes, and abdomen with middorsal row of pale 
triangles; body elongated, especially in females, 
and sometimes with soft abdomens; wings elon- 
gated with clouds accentuated on cross veins, 
first posterior (R;) cells closed and petiolate, 
spur veins present; legs long, particularly the 
fore pair, including fore coxae, hind tibiae with 
two apical spurs; eyes bare, unicolorous (re- 
laxed), those of males widely dichoptic, facets 
undifferentiated in size; three relatively compact 
and elevated ocelli at vertex; fronts of females 
about as wide as high with no basal callosities 
but with some rugosities above subcalli; subcalli 
elongated and with lateral hairs; faces produced 
conically, snoutlike, without denuded areas; 
parafacials narrow; antennae subulate with 8-an- 
nulate flagellums; palpi subcylindrical or flat- 
tened with short hairs and subapical ‘‘sensorial 
grooves’ (containing numerous microscopic or- 
ganelles); proboscides long, basally with unusu- 
al hinge-type articulations, shafts and labella 
strongly sclerotized and extensile; maxillae non- 
serrated. Male interoptic ‘‘fronts’’ narrower 
with longer hairs than in respective females. 

Male genitalia (for terms, refer to Fig. 1): Ab- 
domen abruptly narrowed from segment IV or 
V and bulbous terminally, T. I[X—X shield- 
shaped, not retracted beneath T. VII, as is usu- 
al, and basally straight or convex; cerci project- 
ed distally; branches of gonapophyses (basistyli) 
incompletely sclerotized, membranous on inner 
surfaces, and ventrally with apical styli, plus 
tuberosities on inner bases; flagellar aedeagus 
strongly extended cephalad; apodeme of aede- 
agus spatulate with mid-longitudinal keel. Fe- 
male: T. 1X undivided and with peculiar ‘‘flaps”’ 
projecting distally; T. X frequently undivided; 
genital forks weakly sclerotized, especially ba- 
sally, and with small comb-teeth; caudal ends of 
spermathecal ducts with peculiar, intermediate 
sclerotized parts, continued as long tubes cov- 
ered by what we term ‘‘small alimentary tu- 
bules,”’ and apically, neither bulbous nor scle- 
rotized; St. VIII wide, frequently with basal 
portion very curved and strongly sclerotized lat- 
erally; gonapophyses small, subglobose and well 
sclerotized; no externally visible cerci projected 
over gonapophyses. 

Generalized, nonhaematophagous, flower 
feeders. 


Promycteromyia Coscaron and Philip, 
new genus 
Type-species.—Mycteromyia philippii Philip 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


DIAGNOSIS.—Species grayish to dark grayish 
yellow or blackish. Male: Basistyli with tuber- 
osities inwardly below; dististyli spoon-shaped; 
aedeagus funnel-shaped (dorsal view), gonapo- 
physes basoventrally strongly curved cephalad, 
and well sclerotized basolaterally. Female: T. X 
undivided; St. VIII with a strongly curved, well- 
sclerotized base projected laterally. 

INCLUDED SPECIES.—Promycteromyia philip- 
pii (Philip), P. galbina new species, P. derocer- 
ca new species, P. eriodes (Philip), P. cineras- 
cens (Bigot), P. pechumani new species, P. 
penai new species, P. murina (Philippi), and P. 
xantha new species. 


Promycteromyia philippii (Philip), new combi- 
nation 

(Figure 1) 

Mycteromyia philippii PHILIP, 1958:63. 

Mycteromyia conica: PHILIPPI (not Bigot), 1865:712. 

D1AGNosis.—Medium-large, blackish species, 
abdomen dark brown dorsally, black haired, 
gray pollinose and pale pilose on T. I, on basal 
half of T. Hf, and on mid-stripe of truncated tri- 
angles, as well as often on lateral incisures (Fig. 
1A). Beard pale pilose; ventrally, thorax and ab- 
domen, coxae and femora brownish black with 
black hairs; pleura grayish pollinose, pale 
haired. 

DESCRIPTION.—Male. Lengths 12-13 mm, 
wings 10-11 mm, proboscides 5—8 mm, ratio pal- 
pus:wing 1:7—9. Genitalia: Visible cercus sub- 
triangulate with rounded apex (Fig. 1B), in slide 
view with small external emargination (Fig. 1C); 
style strongly curved with wide base, dististyle 
relatively short with strong emargination on dis- 
tal border and with a strong ridge on proximal 
border (Fig. 1D). 

Female. Lengths 12-15 mm, wings 10.5—13 
mm, proboscides 6.5-8.5 mm, ratio palpus: wing 
1:9.5-10. Genitalia: S. VIII with deep concavity 
basally and gonapophyses strongly sclerotized; 
cercus subtriangulate; T. IX short, with large 
lateral flaps (Fig. IF); sclerotized portions of 
ends of caudal parts of spermathecal ducts with 
‘“‘twists’’ that are continued caudad in the ducts 
(Fig. 1E£). 

MATERIAL EXAMINED (all from CHILE).—Coquimbo: 1 
paratype female, El Sauce, Elqui, 5.XI.1937 (E. P. Reed); 2 
males, | female, El Pangue, 3.XI.1961 (Pena); 1 female, El 
Pangue, 29.X.1937 (Reed); 1 female, El Tongay, Huanaquero, 
28.X1.1952; 15 females, Hacienda IIlapel, El Calabazo, 21- 


22.X1.1961 (Pena); 2 females, Hacienda Illapel, Canela Baja, 
23-24.X.1961 (Pena); 1 female, Socos, 13.X.1957 (Pena), and 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


FIGURE 2. 
dorsum. F. Epandrium and genital appendages. G. Epandrium and cerci, caudal view. H. Epandrium, cerci and ventral plate 
in slide view. J. Basistyli, dististyli and aedeagus. J. Dististyle, dorsal view. 


2 males, 1 female, same data but 16.1X.1963; 1 female, Cuesta 
de los Hornos, 18.X.1958 (Pena); 1 female, 8 km N San Pedro 
de Quile, 16-17.X1.1961 (Pena); 1 female, “‘Entrance to the 
Tunnel Aconcagua,’ 90 km S IIlapel, 28.XI.1950 (Ross and 
Michelbacher). Atacama: 1 female, Quebrada del Algodo- 
nero, 18.X.1957 (Pena). 


Promycteromyia galbina Coscaron and Philip, 
new species 
(Figure 2) 


DIAGNosis.—Medium-sized, grayish-yellow 
species; mid-abdominal stripe of connected tri- 


Promycteromyia galbina new species, male. A. Head. B. Antenna. C. Palpus. D. Head in profile. E. Abdomen, 


angles with wide bases and long pale hairs on T. 
II-VI, enclosed in a wide, brown-haired stripe 
and with longer yellow hairs on sides (Fig. 2E); 
venter gray-brown. 

DEscRIPTION.—Holotype male. Length 13 
mm, wing 10.5 mm, proboscis 8.4 mm, ratio pal- 
pus:wing 1:8.2. Front and face grayish with 
small lateral brown bands; front (Fig. 2A) with 
long black hairs above; scape and pedicel (Fig. 
2B) gray pollinose; palpus (Fig. 2C) with black 
and yellow pile; proboscis (Fig. 2D) dark brown. 


432 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


x 


7 Las 


FIGURE 3. Promycteromyia derocerca new species. Female (A-H), male (J-M). A. Head in profile. B. Antenna. C. Palpus. 
D. Head. E. External genital appendages, lateral view. F. Same, caudal view. G. Genital fork; cd, caudal spermathecal ducts; 
sc, sclerotized portions (twisted). H. Genital appendages. J. Genital appendages, lateral view. J. Same, caudal view. K. 
Epandrium and cerci in slide view. L. Style and dististyle. M. Dististyle, ventral view. 


Notum grayish with four prominent brown _ black distally, tibiae and tarsi yellowish to pale 
stripes and sparse yellow and black hairs; gray- brown with yellow hairs. Wings subhyaline, 
brown, gray pollinose, and yellow pilose ven- cross veins with very faint clouds; spur veins a 
trally. Coxae and basal halves of femora gray, _ little longer than stems. 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


Allotype female. In close agreement with ho- 
lotype, but legs darker and tergites grayer lat- 
erally. Length 13.5 mm, wing 11 mm, proboscis 
7.5 mm, ratio palpus: wing 1:8.7. Genitalia: Cerci 
subtriangulate, T. IX tall, but a little less so than 
in P. derocerca, and flaps more sinuous lat- 
erally; T. X with a median depression, St. VIII 
as in P. eriodes but deeper; genital fork as in P. 
cinerascens but more sclerotized basally; caudal 
spermathecal ducts very elongated, sclerotized 
portions unknown. 

Paratype males. Very close to holotype in or- 
namentation. Lengths 12.5-13 mm, wings 10- 
10.5 mm, proboscides 7.5-8.5 mm, ratio pal- 
pus:wing 1:8-8.8. Genitalia: Visible cerci sub- 
triangulate with rounded apices, emarginate on 
anterior and posterior borders (Figs. 2F and 2G), 
in slide view, cerci are thin distally (Fig. 2H); 
style straight, dististyle with narrow distal por- 
tion elongated (Figs. 2/ and 2/). 


TyPeE-SERIES (all from CHILE).—Holotype male. Coquim- 
bo: Fray Jorge, 4-5.X1.1951 (Pena) (LLP). Allotype female. 
Same data as for holotype. Paratypes. 1 male, Fray Jorge, 
20.X.1966 (Schlinger and Irwin); 1 male, Fray Jorge, 
9.XI.1971 (Pino); 1 male, Fray Jorge, 4-5.X1.1957 (Pena); 1 
male, Tongay, Huanaquero, 28.X1.1952; 2 males, El Tongue, 
13.X.1957; 1 male, Los Vilos, 13.X.1961; 1 male, Los Vilos 
to Illapel, -.X.1965 (Pena). (CAS, MLP, and MNHN). 


COMMENTS.—This species has a peculiar yel- 
low ornamentation that differs from other relat- 
ed species. Pangonia obscuripennis Philippi, 
which Kroeber (1934) included in Mycteromyia, 
resembles galbina in having yellowish abdomi- 
nal bases and pale middorsal triangles. Philippi’s 
failure to include P. obscuripennis in Myctero- 
myia or to mention the closed first posterior cell 
characteristic of Mycteromyia (plus the now- 
missing type), impels us to omit it from this re- 
view. 


Promycteromyia derocerca Coscaron and Philip, 
new species 
(Figure 3) 


DraGnosis.—Medium-sized, yellowish-brown 
species with abdomen yellow to reddish brown 
dorsally, yellow and black haired with brown 
mid-stripe enclosing a row of continuous, gray- 
ish-yellow-haired triangles, venter brown with 
black hairs; palpi relatively long, and male with 
conspicuous, elongated cerci, strongly acumi- 
nate distally. 

DESCRIPTION.—Holotype male. Length 11.5 
mm, wing 9 mm, proboscis, 6.5 mm, ratio pal- 


433 


pus:wing 1:8. Front and face gray with a brown 
band on each side, front with long black hairs 
and some pale-gray ones below, beard pale; an- 
tennae black, scapes and pedicels grayish pol- 
linose; palpi and proboscis brownish black. No- 
tum gray with brown stripes and sparse pale and 
black hairs; scutellum black, gray pollinose, 
pleura and coxae blackish gray, pale haired. Fe- 
mora gray-brown to reddish brown apically, tib- 
iae and tarsi lighter yellowish brown with yellow 
hairs. Wings subhyaline with brown clouds on 
cross veins, spur veins about twice as long as 
stems. Epandrium gray-brown with pale hairs. 

Paratype males. Lengths 11.5—12.5 mm, wings 
8.5-9 mm, proboscides 6-8.5 mm, ratio pal- 
pus:wing 1:6-6.5. Ornamentation in good 
agreement with type; T. I gray laterally, remain- 
der brownish yellow on sides, incisures grayish 
yellow, epandrium gray-brown. Genitalia: Visi- 
ble cerci strongly protruded, acuminate hooklike 
apically (Fig. 3/); in slide view, very elongated, 
more than half length of epandrium shield (Fig. 
3K); styli very curved (Fig. 3L), dististyli elon- 
gated (Figs. 3L, 3 M). 


TyPeE-SERIES (all from CHILE).—Holotype male. Coquim- 
bo: Los Loritos, 21.X.1967 (Pena), CAS Ent. Type No. 12679. 
Paratypes. Same state, 1 male and 2 females, Cuesta de Bue- 
nos Aires, 14.X.1957 (Pena); 1 female, Los Vilos, 26.X.1965; 
1 female, Huanaquero, Tongoy, 28.X.1952. (CAS, MLP and 
LIL} 2). 


COMMENTS.—Several poorly preserved fe- 
male specimens are tentatively included here, 
but none as types. Two are topotypic; they show 
some resemblance, except in genitalia, to P. phi- 
lippii, with which they are also sympatric. They 
have similar brown to black thoraces and ab- 
domens and T. I and II are white pilose. Typical 
P. derocerca, however, are smaller. The speci- 
mens have brown-gray beards, as in P. murina, 
which has been taken near Santiago and Curico, 
whereas our specimens are from distant Co- 
quimbo. In this group we generally have not seen 
a species with such a wide distribution. Lengths 
11-12 mm, wings 9-10 mm, proboscides 6~7 
mm, ratio palpus:wing 1:7.4-8.2. Color of head 
and appendages as in male, with short gray- 
brown to whitish hairs on head and beard; head 
(Figs. 3A and 3D); antenna (Fig. 3B); palpus 
(Fig. 3C), but occasionally (as in males) more 
acuminate distally, saber-shaped. Thorax grayer 
than in male, pleura whitish pilose; legs gray to 
dark brown; tibiae and tarsi mostly with con- 
colorous hairs. Abdomens grayish brown with 


434 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


FIGURE 4. Promycteromyia eriodes (Philip). Male (A, B), female (C-I). A. Head. B. Basistyle, tuberosity, style and dististyle, 
ventral view. C. Head. D. Antenna. E. Palpus. F. Abdomen, dorsum. G. Sclerotized portion of spermathecal duct; di, diverticle. 
H. Sternite VIII (=subgenital plate) and gonapophyses; Do, dorsal view; g, gonapophyses; Ve, ventral view. /. Genital appen- 


dages. 


pale hairs on median triangles and prominently 
on lateral incisures of T. I and II over gray-pol- 
linose integument (not yellow as in males). Ven- 
ters brown with brown and gray hairs. Genitalia: 
Visible cerci prominent, subtriangulate (Figs. 3E 
and 3F), each with an aperture angle of about 
90°; T. IX very tall, with lateral flaps weakly 
sinuous (Fig. 3H). Caudal spermatheca! ducts 
very long; sclerotized portions with several 


twists (Fig. 3G). St. VIII as in P. cinerascens 
but more deeply emarginated basomesially. 


Promycteromyia eriodes (Philip), new combina- 
tion 
(Figure 4) 
Mycteromyia eriodes PHILIP, 1958:66. 

As this species was originally described in suf- 
ficient detail, only a diagnosis, description of the 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


male genitalia, and the presumed female char- 
acters are given here. 

DIAGNosIs.—Medium-large species; male with 
unusually woolly appearance due to the shaggy 
whitish pile over the entire body; presumed fe- 
male less hairy, abdomen with mostly short 
grayish-white hairs and a white middorsal stripe 
bordered by submedian reddish-brown stripes. 

DESCRIPTION.—Male. Lengths 13.5-14 mm, 
wings 10-10.5 mm, proboscides 9 mm, ratio pal- 
pus: wing 1:7.5; frontal aspect of head as in Fig- 
ure 4A. Genitalia: Externally subtriangulate with 
little emargination posteriorly, much as in P. 
galbina; in lateral view, with emargination on 
inner side very similar to that in P. cinerascens. 
Base of style with two emarginations laterally; 
dististyle elongated distally with strong emargin- 
ation posteriorly (Fig. 4B). 

Female. Lengths 13.5 mm, wings 10.5 mm, 
proboscides 9 mm, ratio palpus:wing 1:6.5-8. 
Head and appendages as in male, but with more 
black frontal hairs; antenna (Fig. 4D). Palpi as 
in male, but not as elongated or pointed distally 
(Fig. 4£). Thorax, legs, and abdomen compa- 
rable to male but integumental pilosity more ev- 
ident beneath sparser vestiture (Fig. 4F). Geni- 
talia: St. VIII with a deep median emargination 
basally (Fig. 4H), cerci more subquadrate than 
in P. derocerca. T. [X narrow and flaps strongly 
sinuous laterally (Fig. 4/); caudal portion of 
spermathecal ducts well sclerotized and with 
about two twists (Fig. 4G). 


MATERIAL EXAMINED (all from CHILE).—Valparaiso: Ho- 
lotype and paratypes from Olmueé; | female, *‘S. For’ (Val- 
paraiso?), 9.1X.1921; 1 female, ‘‘A. foz.’’ Santiago: 1 female, 
Penalolen, 9.II.1944 (Pena). The last specimen is more grayish 
than the others. 


Promycteromyia cinerascens (Bigot), new com- 
bination 
(Figure 5A—F) 


Mycteromyia cinerascens BiGor, 1892:610. 


DrtaGNosis.—Medium-sized, grayish-brown 
species, abdomen with a wide middorsal brown, 
black-haired stripe enclosing a row of continu- 
ous grayish, pale-haired triangles, and laterally 
gray on T. I, gray to reddish brown on T. II, 
and blackish gray on sides of remaining tergites, 
covered with yellowish-gray hairs; visible cerci 
trapezoidal-shaped. 

DESCRIPTION.—Male. Lengths 10.5-12 mm, 
wings 8-10 mm, proboscides 6-8 mm, ratio pal- 
pus:wing 1:6-7.5. Front and face gray; front 


435 


with long, mostly black hairs and with a few 
yellow hairs below; face with shorter yellow 
hairs; beard yellow to whitish. Antennae black, 
scapes and pedicels gray pollinose. Palpi gray- 
black with black and yellow hairs. Thorax 
brownish gray with yellow to brown hairs dor- 
sally, and yellowish hairs laterally, blackish 
brown with sparse yellow hairs ventrally; coxae 
and femora blackish brown with yellow hairs; 
tibiae and palpi light reddish brown with yellow 
hairs. Abdomen brownish black with pale inci- 
sures ventrally. Genitalia: Cerci externally as 
figured (Fig. 5A); in slide view, emarginated in- 
wardly (Fig. 5B); styli gently curved, attenuated 
on apical halves, dististyli elongated on fore bor- 
der, slightly emarginated (Fig. SC). 

Female. Lengths 10.5—13 mm, wings 9.5-11.5 
mm, proboscides 5.5-6 mm, ratio palpus: wing 
1:9.3-10. Head and appendages as in male, but 
front (as in related species) wider and shorter 
haired. Hairs of palpi yellowish to black, of 
beard white to yellow, of thorax brownish gray; 
pleura mostly whitish haired; coxae and femora 
grayish black with black and white hairs; tibiae 
and tarsi brownish gray with black pile. Abdo- 
men as in male but lateral pale areas more re- 
stricted and grayish with mostly pale pilosity; 
venter light gray to brown with sparse pale hairs. 
Genitalia: St. VIII concave, base not strongly 
emarginated mesially (Fig. 5E), cerci subtrian- 
gulate, T. IX with very sinuous-margined flaps 
laterally (Fig. 5F); genital fork sclerotized ba- 
solaterally, and with very long, peculiar sper- 
mathecal ducts that are occasionally sclerotized 
on the anterior halves, the ends of the sclero- 
tized portions with one to two twists (Fig. 5D). 

MATERIAL EXAMINED (all from CHILE).—Santiago: 7 
males, 2 females, Aculeo, El Arbol, -.X.1969 (Pena). Valpa- 
raiso: | male, 20 km N Concon, 26.X1.1950 (Ross and Mich- 
elbacher); 1 male, Cuesta de Llay Llay, 20.X.1961 (Pena). 
Valparaiso?: 6 males, El Noviciado, 2-4.X.1954 (Pena). Co- 
quimbo: 6 males, Hacienda Illapel, El Calabazo, 21- 
22.X1.1961 (Pena); 2 males, Canela Baja, 23-24.X.1961 
(Pena): 1 male, 3 females, El Tongue, 13.X.1957 (Pena); 1 
male, 1 female, Chanaral del Aceituno, 23—25.X.1957 (Pena). 
Atacama: 2 females, Vallenar, 28.1X.1957 (Pena). 

COMMENTS.—This species 1s difficult to char- 
acterize because of its variability. Males vary 
from light gray with pale pile and femora brown- 
ish gray with white to darker pile, to reduced 
pale sides on the abdomen, and tibiae and tarsi 
brown with black hairs. In females, we observed 
variation in size, in vestiture more extensively 
pale brown, and in legs paler on tibiae and tarsi; 


436 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


YY \ 
IN \NS 
aM \K m 
ZX) VA WN 
YX | iN 
SHY VK WX 
Lim A 
Ay A) YN 
G7 Ay YN 
wor ey 


FIGURE 5. 


Promycteromyia cinerascens (Bigot). Male (A-C), female (D-F). A. External genital appendages. B. Epandrium 


and cerci. C. Style and dististyle. D. Genital fork and spermathecal ducts. E. Sternite VIII and gonapophyses. F. Genital 
appendages. Promycteromyia pechumani new species, male. G. External genital appendages. H. Epandrium and cerci. J. Style 


and dististyle. 


thus being very difficult to match with males, 
especially as this species is sympatric with sev- 
eral related ones. 

Promycteromyia cinerascens from Olmué, 
Valparaiso, was correctly identified by Kroeber 
(1930a), but Barretto and Duret (1954) mistak- 
enly assigned a male from Rio Negro, Argentina, 
to this species. Their description and genitalic 
drawings differ from our interpretation of P. ci- 
nerascens. The different shapes of cerci and dis- 


tistyli plus the short styli and valve of the ae- 
deagus with strong bristles relate their specimen 
to Silvestriellus and close to our S. schlingeri 
new species. 


Promycteromyia pechumani Coscaron and Phil- 
ip, new species 
(Figure 5G-/) 

DIAGNosIs.—Medium-sized, grayish-brown 
species, homogeneously colored abdomen gray 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 437 


FIGURE 6. Promycteromyia penai new species. Female (A-C), male (D-F). A. Head. B. Head in profile. C. Sclerotized 
portion of spermathecal duct. D. External genital appendages. E. Epandrium and cerci. F. Basistyli and dististyli. Promycter- 
omyia murina (Philippi). Male (GJ), female J). G. External genital appendages. H. Epandrium and cerci. /. Style and dististyle. 
J. Genital appendages. 


dorsally with middorsal stripe of pale triangles black hairs; laterad of triangles on T. II and III 
over obscurely marked integument. Vestiture are black to brown hairs which do not form the 
very sparse, grayish yellow on triangles, and usual continuous, wide brown stripes observed 
flanked by darker, sparse, mixed brown and in most related species. Visible cerci of male 


with small emarginations above and below dis- 
tally (Fig. SG). 

DESCRIPTION.—Holotype male. Length 12.5 
mm, wing 10.5 mm, proboscis 7 mm, ratio pal- 
pus: wing 1:9.5. Front dark gray with black hairs 
above, brownish below and on face. Palpus 
blackish-gray pollinose with black hairs. Thorax 
brownish gray with intermixed black and yellow 
hairs. Pleura, femora and coxae brownish-gray 
pollinose with pale hairs, femora a little lighter 
than coxae, tibiae and tarsi light brown with 
brownish-black hairs. Venter dark gray with pal- 
er incisures and pilosity. 

Allotype female. In good agreement with male 
but without brown pattern. Length 13 mm, wing 
11 mm, proboscis 8.5 mm, ratio palpus:wing 
1:8.5. Genitalia very close to P. cinerascens but 
with St. VIII less sclerotized, cerci more 
subquadrate, and flaps of T. IX more sinuous 
laterally; sclerotized portion of caudal sper- 
mathecal ducts lightly sclerotized. 

Paratype males. Agree with holotype although 
abdominal pattern sometimes more evident. 
Lengths 11.5-12.5 mm, wings 9-10 mm, probos- 
cides ca. 8 mm, ratio palpus: wing 1:9. Genitalia: 
Cerci in slide view with reduced emarginations 
laterally, acuminate, but rounded apically (Fig. 
5H), styli thin and curved, dististyli with strong 
ridge anteriorly and strongly emarginated pos- 
teriorly (Fig. S/). 


TyPe-SERIESs (all from CHILE).—Holotype male. Aconca- 
gua: Valle de Piuquenes, 14~-25.XI.1958 (Pena) (LLP:. Allo- 
type female. Same data as holotype. Paratypes. Aconcagua: 
1 male, topotypic; 1 male, Guardia Vieja, 8.XII.1958 (Pena); 
(CAS and MLP). 


COMMENTS.—This species can be differen- 
tiated from P. derocerca, P. cinerascens, and P. 
murina by its homogeneous gray color, and from 
P. penai new species by lacking coarse pile and 
by having wide cerci with two subapical emar- 
ginations. This species is dedicated to L. L. Pe- 
chuman who provided much of the material uti- 
lized in this study. 


Promycteromyia penai Coscaron and Philip, new 
species 
(Figure 6A-F) 


DIAGNosIs.—Small, gray-colored species with 
sparse but unusually erect pale hairs, second 
segments of palpi shorter than antennal flagella 
(Fig. 6B), visible cerci rounded or truncated in 
males. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


DESCRIPTION.—Holotype male. Length 10 
mm, wing 9 mm, proboscis 5.5 mm, ratio pal- 
pus:wing 1:11. Front gray with mostly yellow 
hairs, some black ones above; subcallus and 
face pale gray with yellow hairs. Thorax, coxae, 
and femora brownish gray with pale hairs; tibiae 
and palpi yellowish brown with whitish hairs. 
Abdomen gray with double rows of paired, sub- 
median, discontinuous brown spots, narrowed 
caudad. Venter gray with pale pile. 

Allotype female. Agrees well with holotype; 
length 10.5 mm, wing 9 mm, proboscis 5.5 mm, 
ratio palpus:wing 1:9. 

Paratypes. Males agree with the holotype. 
Male genitalia: Visible cerci a little longer than 
wide with distal margin rounded or truncated 
(Fig. 6D); in slide view, dorsobasally narrowed 
distally and ventrally (Fig. 6£); styli thin and 
curved, dististyli strongly emarginated, thin and 
elongated distally (Fig. 6F). Female also with 
similar ornamentation. Head as in Figure 6A. 
Female genitalia: St. VIII as in P. cinerascens, 
but with a median emargination basally and less 
sclerotized laterally, cerci subtriangulate, T. LX 
shorter then in P. galbina, and flaps not very 
sinuous laterally, T. X weakly sclerotized with 
only one diverticle (Fig. 6C). 


TyYPE-SERIES (all from CHILE).—Holotype male. Coquim- 
bo: Choros Bajos, 31.X.1961 (Pena), CAS Ent. Type No. 
12680. Allotype female. Same data as for holotype. Paratypes. 
1 male, Los Loritos, 26.X.1957 (Pena); 1 male, 2 females, 
Chanaral del Aceituno, 15—23.X.1957; (CAS, LLP and MLP). 


COMMENTS.—Coloration very similar to P. 
pechumani new species except lacking yellow 
tones, but the smaller size, greater hairiness, 
shorter palpi, and different external shapes of 
cerci, permit differentiation. This species is ded- 
icated to Luis E. Pena, our Chilean entomolog- 
ical friend, who originally collected most of our 
study material, much of which was loaned 
through the courtesy of Dr. L. L. Pechuman. 


Promycteromyia murina (Philippi), new combi- 
nation 
(Figure 6G—J/) 


Mycteromyia murina PHILIPPI, 1865:713. 


Philip (1968) saw and redescribed the pre- 
sumed male type of this species in the Museo 
Nacional de Historia Natural in Santiago. It 
lacked the original author’s labels and was se- 
lected from other similarly unlabelled ‘‘Types”’ 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


by Kuschel from among the Philippi specimens 
and could therefore be questionable. This 
‘‘type,’’ according to Philip (1968:12), has a pe- 
culiar ‘‘dark brown beard, chest hairs and fe- 
mora,’ ‘‘abdomen, including the venter is dull 
black,’’ and “‘visible cerci rounded and coarsely 
black-haired, not pointed apically as in ciner- 
ascens.’’ Among specimens known to us which 
fit this description and with rounded cerci, we 
have two candidate groups: one group of spec- 
imens from the Coquimbo area, with a smaller, 
grayish-hirsute body, and the other from Santia- 
go and Curico with a larger, brown, and less 
hirsute body. We assign, as first revisors, the 
second group to murina. 


DIAGNosIsS.—Medium-sized species with 
blackish to brownish-gray abdomens, and wide, 
distally rounded or truncated cerci in males. 


DESCRIPTION.—Male. Lengths 12.5—13.5 mm, 
wings 8-10 mm, proboscides 8-9 mm, ratio pal- 
pus:wing 1:7—-7.4. Front gray with black hairs 
above to yellowish brown below, beard varying 
from black to mixed brown and yellowish gray; 
distal article of palpus a little longer than anten- 
nal flagellum; thorax brownish gray, notal hairs 
brown to yellow and pleura pale; coxae and fe- 
mora dark with black to grayish-yellow hairs, 
tibiae and palpi reddish brown with mixed black 
and gray pilosity. Abdominal integument gray to 
brownish yellow dorsally with paired brownish 
submedian dashes margining gray median tri- 
angles. Tergite I and epandrium brownish gray. 
Abdominal vestiture pale dorsally. Venter most- 
ly blackish brown with black hairs and paler in- 
cisures. Genitalia: Visible cerci approximately 
as wide as high, with rounded or truncated mar- 
gins (Fig. 6G): in slide view, subquadrate, and 
truncated distally (Fig. 6H). Styli relatively 
shorter than in P. penai, approximately as wide 
as high; dististyli elongated, weakly emarginated 
caudad and cephalad (Fig. 6/). 

Female. Lengths 13.5-14 mm, wings 12-13 
mm, proboscides 8-8.5 mm, ratio palpus: wing 
1:7.8-8.4. Color resembles that in the male with 
similar variations, but abdominal sides mostly 
yellow. Genitalia: St. VIII with shape as in P. 
cinerascens , but with smaller median concavity, 
and less sclerotization laterally. Cerci subquad- 
rate, T. [X wide, nonsinuous laterally and with 
a peculiar transverse, subbasal, sclerotized bar 
(Fig. 6/); genital fork with a moderately scle- 
rotized base; caudal spermathecal ducts elon- 


439 


gated, with sclerotized portions not twisted, and 
with only one lateral diverticle. 


MATERIAL EXAMINED (all from CHILE).—Santiago: 1 
male, Maipu, Quebrada de La Plata, 11.X.1966 (Schlinger and 
Irwin); Curico: 1 male, El Coigo, -.II.1961 (Pena); 1 male, El 
Buchen, 12.11.1961 (Pena); 1 male, 1 female, Teno River, 19- 
20.1.1964 (Pena); | male, Vergara River, 2,000-2,300 m, 
26.1.1968 (Pena); 1 male, 1 female, La Vinilla, 19.11.1961 
(Pena); 3 males, Teno River, 1,800 m, -.I1.1965 (Pena). 


Promycteromyia xantha Coscaron and Philip, 
new species 
(Figure 7A-G) 


DIAGNosIs.—Large, soft-bodied, distinctive, 
light brownish-yellow species, abdomen mostly 
dark haired except on paler outer incisures, with 
a brown mid-stripe enclosing a row of discontin- 
uous, pale triangles. 

DESCRIPTION.—Holotype male. Length 15.5 
mm, wing 12.5 mm, proboscis 7 mm, ratio pal- 
pus:wing 1:11.3. Front, scape, pedicel, and pal- 
pus yellowish-gray pollinose, frontal pilosity 
mostly yellow with some darker hairs inter- 
mixed; face and subcallus pale yellow. Scape, 
pedicel, and palpus with black pile; flagellum 
missing. Front as figured (Fig. 7A). Head profile 
as in Figure 7B; palpus relatively short (Fig. 7C). 
Beard and pleura with pale hairs. Notum with 
brown stripes; scutellum lighter on disk than in 
P. murina, with yellowish and a few black hairs. 
Wing veins light brown, membrane opalescent. 
Legs light brown with pale pollinosity; coxae 
and femora with yellow hairs, while the tibiae 
and tarsi have dark ones. Mid-abdominal row of 
pale triangles, T. I and all incisures with yellow 
pile (Fig. 7D). Venter yellowish brown with pale 
hairs except dark ones on hind margins of St. 
II-VI. Genitalia: Visible cerci relatively short, 
narrow, and rounded distally; in slide view, 
short, curved mesoventrally and rounded dis- 
tally (Fig. 7G); epandrium more flexuous than 
usual. Styli very curved and thick, dististyli 
elongated without emargination cephalad or 
caudad (Figs. 7E and 7F). 


TypE-Data.—Holotype male. CHILE, Talca: Alto de 
Vilches, 26-27.1.1964 (Pena), CAS Ent. Type No. 12681. 


COMMENTS.—This is a peculiar species, un- 
fortunately only represented by one somewhat 
damaged but unusual specimen. It can be distin- 
guished by its coloration from all other species 
of the Promycteromyia group that have distinc- 
tive tuberosities on the inner sides of the gona- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


440 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


pophyses. Caenopangonia brevirostris (Philip- 
pi), which has similar coloration and a soft 
abdomen, is readily distinguished by its short 
face. 


Genus Mycteromyia Philippi 


Mycteromyia PHILIPPI, 1865:712. [Type-species: Pangonia 
conica Bigot.] 


DESCRIPTION.—Large brown species with 
long palpi and strongly clouded and tinted 
wings; male basistyle without tuberosities ven- 
tromesially, dististyle subtriangulate without 
heavy midventral ridge, and with subapical pro- 
tuberance on anteroventral borders; styli long 
and straight; aedeagus wide distally with lateral 
valves smooth; gonapophyses with ventral base 
slightly convex and not sclerotized laterally. Fe- 
male T. X divided; St. VIII wider than high, not 
strongly sclerotized laterally; genital fork with- 
out sclerotized base, caudal spermathecal ducts 
with lateral diverticles. 

INCLUDED SPECIES.—Mycteromyia conica 
(Bigot) and M. etcheverryae new species. 


Mycteromyia conica (Bigot) 
(Figure 8) 


Pangonia conica BiGoT, 1858:278. 
Mycteromyia fusca PHILIPPI, 1865:712. 


DIAGNosIs.—Large brown species with mid- 
abdominal row of pale triangles, wings generally 
brown and with darker clouds on cross veins 
(Fig. 8C), palpi elongated, saber-shaped. 

DESCRIPTION.—Male. Lengths 15-17 mm, 
wings 14-15 mm, proboscides 10.5—11 mm, ratio 
palpus:wing 1:7. Female. Lengths 19-20 mm, 
wings 15.5-17 mm, proboscides 12 mm, ratio 
palpus:wing 1:7.5—8.5. Front gray pollinose, 
generally with wide, darker stripe from ocellar 
tubercle to bases of antennae; face produced 
snoutlike (Figs. 8A and 8B), pollinose with wide 
brown band on each side. Frontal hairs blackish 
brown, sometimes with a few yellow ones. 
Scape and pedicel brownish-gray pollinose with 
black hairs; flagellum reddish brown. Palpus 
with black hairs. Beard brownish. Notum with 


44] 


prominent brown bands and black to pale 
haired, pleura brownish-gray pollinose with pale 
to brownish-black pilosity. Basicosta nonsetu- 
lose (Fig. 8D). Legs reddish brown to black with 
pale hairs on coxae and femora, and golden to 
dark ones on tibiae and tarsi. Abdomen dorsally 
light to dark brown and dark haired, with median 
stripe of continuous elongated triangles, and 
sometimes pale lateral incisures; venter dark 
brown. Male genitalia: Visible cerci in lateral 
view triangulate (Fig. 8F), in posterior view with 
apical hooks crossed (Fig. 8£); in slide view, 
subcylindrical (Fig. 8H). Style and dististyle as 
figured (Fig. 8G); dististyle with subapical pro- 
tuberance on anteroventral border, and weakly 
bifurcated apically. Female genitalia: St. VIII as 
figured (Fig. 8M); cerci subquadrate (Fig. 8L). 
Genital fork as figured (Fig. 8/); caudal portions 
of spermathecal ducts well sclerotized with only 
one diverticle each (Fig. 8K). 


MATERIAL EXAMINED (all from CHILE).—Valparaiso: 2 
females, slope of Campana mountain, 1,000 m, 17.XII.1950 
(Ross and Michelbacher). Santiago: 1 female, El Manzano, 
-.XI.1951 (Pena); 1 male, El Quisco, 1-3.XI.1951 (Pena); 1 
male, 1 female, La Obra, -.XII.1951 (Etcheverry); 1 male, 1 
female, El Canelo, 3.XII.1951 (Etcheverry); 5 males, 1 female, 
El Canelo, -.XI.1951 (Etcheverry); 4 males, 1 female, 
-.XII.1951 (Etcheverry); | female, El Canelo, -.XII. 1950 (Etch- 
everry); 4 males, El Canelo, -.XII.1936; 1 female, El Canelo, 
-.X1.1951 (Etcheverry); | female, Penalolén, 8.1.1964 (Mon- 
tero). Colchagua: | male, Nancagua, -.II.1944 (Pena). Curico: 
1 female, Teno River, 6 km E Los Quenes, 5.I.1967 (Irwin). 
Malleco: 1 male, Onate, 29.XI.1958. O'Higgins: 1 female, 
Pangal, 3.1.1964 (Etcheverry); 1 female, Machali, 4.XII.1965 
(Silva). Cautin: 1 male, Tolten, -.I.1955 (Toro). 


COMMENTS.—This species shows much color 
variation, with the body and appendages varying 
from dark brown with brown wings to lighter 
brown with pale outer tergal corners. Myctero- 
myia fusca Philippi was redescribed by Philip 
(1958) and found to be misdetermined conica 
(Bigot). Philip also redescribed M. conica sensu 
Philippi, not Bigot, as M. philippii Philip. Mack- 
erras (1955) made a detailed redescription, in- 
cluding the genitalia, which he considered as so 
distinctive ‘‘as almost to justify removing this 
genus to a separate tribe.” 


<< 


FIGURE 7. 


Promycteromyia xantha new species, male. A. Head. B. Head in profile. C. Palpus. D. Abdomen, dorsum. E. 


Basistyli, dististyli and aedeagus. F. Dististyle and tuberosity. G. Apical portion of epandrium and cerci. Mycteromyia etch- 
everryae new species. Male (H—J), female (K, L), H. Genital appendages, caudal view. J. Same, lateral view. J. Epandrium 
and cerci. K. Genital fork and caudal spermathecal ducts. L. Genital appendages. 


442 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


FIGURE 8. Mycteromyia conica (Bigot). Male (A—H), female (I-M). A. Head. B. Head in profile. C. Wing. D. Basicosta. E. 
Genital appendages, caudal view. F. Same, lateral view. G. Gonapophyses, aedeagus and dististyli. H. Epandrium and cerci. 


I. Head in profile. J. Genital fork and spermathecal ducts. K. Sclerotized portion of spermathecal ducts. L. Genital appendages. 
M. Sternite VIII and gonapophyses. 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


Mycteromyia etcheverryae Coscaron and Philip, 
new species 
(Figure 7H-L) 


DIAGNosIs.—Large brown species, very close 
to M. conica (Bigot) but males with shorter cer- 
ci, and females with caudal spermathecal ducts 
shorter with short nonsclerotized diverticles 
cephalad as shown in respective figures. 

DESCRIPTION.—Holotype male. Length 17 
mm, wing 16 mm, proboscis 10 mm, ratio pal- 
pus:wing 1:9. Front and face gray pollinose, tri- 
ocellar tubercle raised in mid-vertex, and sides 
of snoutlike face brown. Antennae, palpi, and 
proboscis dark brown. Thorax brown, dorsally 
with grayish and pale-brown stripes. Beard, 
pleura, and coxae I with pale pilosity. Wings as 
in M. conica. Legs dark brown with black hairs, 
intermixed with longer pale ones on coxae and 
femora dorsally. Abdomen dark brown with a 
middorsal stripe of white, elongated triangles; 
outer tergal incisures white haired. 

Allotype female. Resembles male but reddish 
brown with thorax, legs, and abdomen lighter 
than in type. Length 15.5 mm, wing 13.5 mm, 
proboscis 8.5 mm, ratio palpus:wing 1:9. Geni- 
talia: St. VIII as in M. conica; cerci rounded, T. 
IX short with wide laterally sinuose flaps (Fig. 
7L); genital fork not sclerotized basally but with 
peculiar, relatively short, robust, unsclerotized, 
caudal spermathecal ducts and small diverticles 
on the sclerotized cephalad portion (Fig. 7K). 

Paratypes. Males are very similar in ornamen- 
tation to the holotype; lengths 15-16 mm, wings 
14-17 mm, proboscides 8-17 mm, ratio 
palpus:wing 1:8-9. Genitalia: Visible cerci sub- 
trapezoidal (Fig. 7/), shorter than in M. conica 
as shown in posterior view (Fig. 7H); in ventral 
view, subtriangular, with evident internal hooks 
(Fig. 7/); gonapophyses very similar to those of 
M. conica, but style of basistyle longer and 
apodeme of aedeagus wider than in conica. 


TypE-SERIES (all from CHILE).—Holotype male. Valpa- 
raiso: Quebrada San Jeronimo, 7.XI.1965 (Arrau) (CEEUC). 
Allotype female. Valparaiso: Quebrada San Jeronimo, 
26.X.1965 (Montes) (CEECU). Paratypes: 4 males, same lo- 
cality and date as holotype, but collected by Zapata; 1 male, 
same locality and date as holotype, but collected by Herrera. 
Santiago: | male, 1 female, El Canelillo, 17.X.1965 (Etchev- 
erry). 


COMMENTS.—This species resembles M. con- 
ica in Ornamentation and size, but it is readily 
separated by differences as given in the key. We 


443 


dedicate this species to Maria Etcheverry, en- 
thusiastic Chilean entomologist, who loaned us 
much Mycteromyia material. 


Genus Silvestriellus Brethes 


Silvestriellus BRETHES, 1910:473. [Type-species: Silvestriellus 
patagonicus Brethes.] 


DESCRIPTION.—Medium to small species with 
relatively short palpi and wing subhyaline with 
basicosta bare; male basistyli without tuberosi- 
ties and styli very short, cerci without apical 
hooks, dististyle without heavy ridge or subapi- 
cal protuberances on anteroventral borders, ae- 
deagus thin distally with decurved hooks at sides 
of valves. Female genital fork well sclerotized 
basally; caudal ends of spermathecal ducts with- 
out lateral diverticles; St. VIII about as wide as 
high, not strongly sclerotized laterally. 

INCLUDED SPECIES.—Silvestriellus patagoni- 
cus Brethes, S. schlingeri new species, S. mar- 
tinezi (Barretto and Duret), and S. flaviventris 
(Barretto and Duret). 


Silvestriellus patagonicus Brethes 
(Figure 9E-G) 
Silvestriellus patagonicus BRETHES, 1910:473; BRETHES 

192122: 

DIAGNosis.—Relatively large, reddish-brown 
species with yellowish pollinosity on thorax and 
appendages, wings subhyaline and with very 
faint clouds on cross veins. Head profile, front, 
and palpus as figured (Figs. 9E, 9F and 9G). 
Length 15 mm, wing 11 mm. 


MATERIAL EXAMINED (all from ARGENTINA).—Santa 
Cruz: Holotype, female?, near Santa Cruz, -.XII.1890 
(MACN). Chubut: 1 female?, Puerto Madryn, -.XI.1932 (Pas- 
chetto) (MZSP). 


CoMMENTS.—Unfortunately, the type is bad- 
ly damaged; missing are antennal flagella, one 
palpus, and almost the entire abdomen, and in 
the specimen from Chubut, abdominal segments 
3-10 are missing. Small median grayish-polli- 
nose triangles occur on T. I and T. II. The sex 
is uncertain in both specimens. Wing 11.5 mm, 
proboscis 8.5 mm, ratio palpus:wing 1:20. The 
relatively short palpi (Figs. 9E and 9G), wide 
front (Fig. 9F), and brownish coloration, plus 
probably distinctive genitalic characters of pre- 
sumed related Argentinian species, impel us to 
revive this genus for a distinctive group in the 
new tribe Mycteromyiini. These flies are so far 
known only from Argentina. 


444 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 


FiGurE 9. Silvestriellus flaviventris (Barretto and Duret), male. A. Head in profile. B. Head. C. Palpus. D. Abdomen, 
dorsum. Silvestriellus patagonicus Bréthes, female? E. Head in profile. F. Head. G. Palpus. Silvestriellus schlingeri new 
species. Male (H—J), female (K-M). H. Gonapophyses, aedeagus and dististyli. J. Epandrium and cerci. J. Head in profile. K. 


Genital fork and spermathecal ducts. L. Genital appendages. M. Sternite VIII and gonapophyses. Silvestriellus martinezi 
(Barretto and Duret), female. N. Head. O. Head in profile. P. Palpus. 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


Silvestriellus schlingeri Coscaron and Philip, new 
species 
(Figure 9H—-M) 


DIAGNOSIS.—Small- to medium-sized, dark 
brown species with mid-abdominal row of dis- 
continuous triangles and lateral incisures pale 
haired; palpus II about one-half the length of the 
antennal flagellum. 

DESCRIPTION.—Holotype male. Length 11 
mm, wing 9 mm, proboscis 4 mm, ratio pal- 
pus:wing 1:18. Front, face, scape, pedicel, and 
palpus gray pollinose, ocellar tubercle brown, 
flagellum blackish brown; hairs of front grading 
from black above to grayish yellow below, ex- 
tending onto face. Palpi similar to those of S. 
martinezi, but a little shorter and more acumi- 
nate apically; palpal hairs black, beard white; 
thorax and legs gray pollinose; hairs on meso- 
notum brownish gray, erect, paler on pleura, 
coxae and femora, mostly blackish on tibiae and 
tarsi. Wings subhyaline, veins dark brown, and 
weakly clouded on cross veins. Abdomen gray 
ventrally with paler incisures and vestiture. 
Genitalia: Cerci in slide view without base pro- 
jected into epandrium (Fig. 9/); dististyli weakly 
concave caudad (more flattened, without the 
typical spoon-shape shown in previous genera) 
(Fig. 9H). 

Females: In agreement with males but are 
more grayish; lengths 11.5-12 mm, wings 9-9.5 
mm, proboscides 5—5.5 mm, ratio palpus: wing 
1:22-24. Profile of head (Fig. 9/) a little less con- 
vex than in holotype; abdomens more elongated. 
Genitalia: Cerci subovoid, hypoproct (St. X—XI) 
short and wide, T. [IX with flaps smooth laterally 
(not sinuous) (Fig. 9L); caudal spermathecal 
ducts robust, with reduced sclerotization (Fig. 
9K); St. VIII relatively narrow basally and gon- 
apophyses in middle (Fig. 9M). 


TyPE-SERIES (all from ARGENTINA).—Holotype male, 
allotype female, and 2 female paratypes. Chubut: 20 km N 
Malaspina, 500 m, 13.XII.1966 (Schlinger and Irwin) (MNHN, 
paratypes in CAS and MLP). Neuquén: 2 male paratypes, Las 
Plumas, 450 m, 17.X.1974 (Gentili) (MLP); 1 female paratype, 
S. M. de los Andes, 18.II.1958 (Foerster) (LLP). 


COMMENTS.—This species resembles $. mar- 
tinezi, but the brownish wing, absence of median 
triangles, and the head in profile more convex 
distinguish martinezi. Silvestriellus patagonicus 
is larger and more reddish brown; S. flaviventris 
has mostly yellowish-orange abdomens, and 
cerci basally more elongated. This species is 


445 


named for Dr. E. I. Schlinger of the University 
of California, Berkeley, who generously provid- 
ed most of the material for study. 


Silvestriellus martinezi (Barretto and Duret), 
new combination 
(Figure 9N—P) 


Mycteromyia martinezi BARRETTO AND DuRET, 1954:209. 


DIAGNosiIs.—Medium-sized, grayish-brown 
species without the usual mid-abdominal trian- 
gles, wing with brownish-margined veins. Pal- 
pus robust (Fig. 9P), shorter than antennal fla- 
gellum (Fig. 90). Head as figured (Fig. 9N). 
Lengths 12-12.5 mm, wing 10.5 mm, ratio pal- 
pus: wing 1:17. 


MATERIAL EXAMINED (all from ARGENTINA).—Buenos 
Aires: Holotype and 1 female paratype, Estancia Barrau, 
-.X1I.1946 (Martinez) (MZSP). 


Silvestriellus flaviventris (Barretto and Duret), 
new combination 
(Figure 9A—D) 


Mycteromyia flaviventris BARRETTO AND DuReET, 1954:210. 


D1iaAGNosis.—Relatively small species with 
peculiar yellowish-orange abdomen and median 
dark stripe not well defined nor enclosing pale 
triangles, wing with brownish-margined veins 
and accentuated spur vein. Cerci elongated ba- 
sally into epandrium. Front, profile of head, pal- 
pus, and abdomen as figured (Fig. 9A—D). 
Length 9 mm, wing 8 mm, proboscis 4.5 mm, 
ratio palpus: wing 1:11.5. 


MATERIAL EXAMINED (from ARGENTINA).—Buenos 
Aires: Holotype male, Estancia Barrau, -.XI.1946 (Martinez) 
(MZSP). 


COMMENTS.—The removed genitalia of this 
type have unfortunately become lost. However, 
the describers’ drawings permit assignment to 
the revived genus Silvestriellus. This species 
closely resembles S. schlingeri, but is differen- 
tiated as above. 


Silvestriellus sp. 


Barretto and Duret (1954:204) described a 
small brownish male with long spur vein from 
Gral. Roca, Rio Negro, Argentina, as Myctero- 
myia cinerascens (Bigot). The descriptions and 
figures disagree with our interpretation of ciner- 
ascens and other known species. The genitalic 
drawings indicate that the species should prob- 
ably be included in Silvestriellus. 


446 


The incomplete ornamentation, as revealed in 
the original description does not match that of 
any known species, and the specimen is too bad- 
ly broken, with critical parts missing, to permit 
further characterization or confident assign- 
ment. 


TRIBE SCIONINI 


Genus Caenopangonia Kroeber 


Caenopangonia KROEBER, 1930b:211. [Type-species: Diato- 
mineura hirtipalpis Bigot.] 


DESCRIPTION.—Species medium to large in 
size, yellow to dark brown in color; abdomen 
elongated with fleshy or rigid integument, wings 
elongated with clouds on cross veins, first pos- 
terior (R;) cells closed and petiolate; spur veins 
present; eyes bare, unicolorous; three ocelli on 
vertex; eyes in male dichoptic and facets not 
enlarged above. Female: Front approximately as 
wide as high and convex in profile, without cal- 
lus and no evident transverse sulcus between 
lower front and subcallus; face very short, gen- 
erally with a depression above base of probos- 
cis, which is hinged and long, with fleshy, well- 
tracheated labella; antennae subulate, flagella 6— 
8 annulate; palpi cylindrical or subclavate with 
basal articles relatively long, second with long 
hairs and deep sensorial grooves apically, and 
containing only a few ‘‘sensorial microscopic 
organelles’’; maxillae nonserrated. 

Male genitalia with long flagella and aedeagus 
with round to elongate apodeme; dististyli sub- 
cylindrical, apically rounded or truncate in form 
of two small lobes; female genitalia similar to 
other Pangoniini with distal spermathecal am- 
pullae not sclerotized, bulbous, and caudal ducts 
of spermathecae sometimes very wide. 

INCLUDED SPECIES.—Caenopangonia hirti- 
palpis (Bigot), C. brevirostris (Philippi) and C. 
asper (Philip). 


Caenopangonia hirtipalpis (Bigot) 

(Figure 10A—E) 

Diatomineura hirtipalpis BiGoT, 1892:619. 

Caenopangonia hirtipalpis (Bigot), KROEBER 1930b:221. 

Mycteromyia edwardsi KROEBER, 1930a:131. 

Mycteromyia bejaranoi BARRETTO AND DwuReET, 1954:207, 
new synonymy. 


DIAGNOsIS.—A variable, medium-sized, dark 
brown species with a very short proboscis (Fig. 
10A), relatively wide, parallel-sided front ele- 
vated basally; fumose wings darkened on cross 
veins; abdomen brown with dark hairs and mid- 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


row of pale triangles mostly not crossing tergites 
adorned with silky whitish hairs, and the latter 
also on lateral incisures. 

DESCRIPTION.—Male. Lengths 9-13 mm, 
wings 9-12 mm, proboscides 4-5 mm, ratio pal- 
pus:wing 1:11-19.5. Genitalia: Visible cerci 
rounded; posterior view (Fig. 10B); in slide 
view, elongated, but less so than short portion 
of epandrium (Fig. 10C). Basistyli (Fig. 10D) 
without lateral expansions, dististyli truncated 
distally with small mesial notch (Fig. 102). 

Female. The only specimen available is a little 
lighter brown with triangles and lateral incisures 
well defined, the former crossing tergites. 
Length 12 mm, wing 11 mm, proboscis 4.5 mm, 
ratio palpus:wing 1:13.7. Genitalia: No signifi- 
cant differences from C. brevirostris. 


MATERIAL EXAMINED.—CHILE. Osorno: | male, Pucatri- 
hue, -.I1.1966. Cautin: 1 male, 20 km E Temuco, 8.1.1951 
(Ross and Michelbacher). Arauco: 1 male, Tres Picos, 
6.11.1967 (Casal); 1 male, Pichinahuel, Cord. Nahuelbuta, 1- 
15.1.1959 (Pena); Pillim Pili, 15.1.1954 (Pena). Malleco: 1 
male, Lonquimay, 13.1.1966; 1 male, Lonquimay, Marimen- 
uco, 1.1.1968 (Pena). Concepcion: 1 male, Parque Bot. Hual- 
pen, 10-15.11.1970. Nuble: 1 male, 1 female, Las Cabras, 6— 
31.1.1963 (Pena); 3 males, Las Trancas, 23—29.1.1967 (Pena); 
1 male, Las Trancas, -.II.1969 (Pena); 1 male, Las Trancas, 
-.1.1965 (Herrera); 1 male, Las Trancas, 28.1.1967 (Schlinger). 
Curico: 6 males, El Coigual, 20-26.1.1964 (Pena); 1 male, El 
Coigual, 11.1.1956 (Pena); El Buchen, 8-9.1.1961. ARGEN- 
TINA. Neuquén: Holotype male of Mycteromyia bejaranoi, 
Pucara, S. M. de los Andes, -.XII.1948 (Schajovskoy); 3 male 
paratypes of M. bejaranoi, same data as holotype but col- 
lected 22.XII.1950, -.X1.1952, and 5.1.1951. 


Caenopangonia brevirostris (Philippi), new com- 
bination 
(Figure 10F—J) 


Mycteromyia brevirostris PHILIPPI, 1865:713. 


D1aGNosis.—The type is lost but the original 
description permits the identification of our ma- 
terial. Medium to large, reddish-brown species 
without prominent abdominal pattern, with no 
or only small median triangles, and very short 
face, as discussed by Philip (1968) and Kroeber 
(1930a). 

DESCRIPTION.—Female. Lengths 11.5-14 mm, 
wings 11-12.5 mm, proboscides 4-4.5 mm, ratio 
palpus: wing 1:8-13. Body and appendages most- 
ly light brownish-gray pollinose with black hairs 
on head and its appendages, pleura, and legs; 
venter pale haired. Well-preserved specimens 
have mid-abdominal rows of pale-yellow trian- 
gles and narrow, lateral, pale incisures. Head 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 447 


Figure 10. Caenopangonia hirtipalpis (Bigot), male. A. Head. B. External genital appendages, caudal view. C. Epandrium, 
cerci and ventral plate of proctiger. D. Gonapophyses, aedeagus and dististyli. E. Dististyle. Caenopangonia brevirostris 
(Philippi), female. F. Head in profile. G. Palpus. H. Sternite VIII and gonapophyses. /. Genital fork and spermathecal ducts. 
J. Genital appendages. Caenopangonia asper (Philip), male. K. Head. L. Palpus. M. Head in profile. N. Gonapophyses, 
aedeagus and dististyli. 


448 


profile and palpus as figured (Figs. 10F and 
10G). Genitalia: St. VIII and gonapophyses well 
sclerotized basally (Fig. 10H); genital fork with 
short caudal spermathecal ducts and abundant 
spines on comb (Fig. 10/); cerci subtriangulate, 
T. X very narrow and T. IX wide, laterally ex- 
panded (Fig. 10/). 

MATERIAL EXAMINED (all from CHILE).—1 female, Chile, 
‘‘Alte Sammlung’? (NMW). Osorno: 3 females, Pucatrihue, 
-.1.1968 (Pena). Arauco: 1 female, Caramavida, 30.1.1967 
(Pena); 1 female, Pillim Pili, 2-5.11.1959; 3 females, Nahuel- 
buta, 20 km W Caramavida, 30.1.1967 (Schlinger). Pichina- 
huel: 1 female, Cord. Nahuelbuta, 28.1.1954. No males avail- 
able. 

COMMENTS.—The variation in size and orna- 
mentation of our inadequate series of speci- 
mens, some of them with variable median tri- 
angles, suggests that as more material becomes 
available, this species may be synonymized with 
the sympatric and also widely variable hirtipal- 


pis. 


Caenopangonia asper (Philip), new combination 
(Figure 10K—N) 


Mycteromyia asper PHILIP, 1958:65. 


DIAGNosIs.—Small- to medium-sized, dark 
brown species having broad yellow, lateral ab- 
dominal incisures and mid-row of continuous 
pale triangles with wide bases. Face in profile, 
moderately produced (Fig. 10K), palpus II rel- 
atively thin and short (Fig. 10L), antenna with 
basal segments and flagellum more slender than 
usual. 

DESCRIPTION.—Male. Length 10 mm, wing 
9.5 mm, proboscis 4 mm, ratio palpus:wing 
1:13.2. Genitalia: Cerci resemble those of C. hir- 
tipalpis but more elongated, about as long as 
mid-longitudinal length of epandrium; gonapo- 
physes laterally expanded, and dististyli acu- 
minate but with blunt apexes (Fig. 10N). 

Female. We now have previously unknown 
females that can be associated readily with 
males. Lengths 10.5—12 mm, wings 10-11.5 mm, 
proboscides about 4 mm, ratio palpus:wing 
1:16-21. Genitalia: Compared with females of C. 
brevirostris, St. VIII has a narrower base, gon- 
apophyses more robust with deeper mesial 
emarginations, and T. IX larger; genital forks 
with bases not mesially sclerotized; distal por- 
tions of spermathecae subovoid, relatively large 
and lightly sclerotized. 


MATERIAL EXAMINED (all from CHILE).—Coquimbo: 1 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


male, 2 females, Fray Jorge, 4-5.XI.1957 (Pena); 1 female, 
Fray Jorge, 9.X.1971 (Pino). 


COMMENTS.—The pointed dististyli and the 
laterally expanded gonapophyses in males, bul- 
bous and distally expanded spermathecae in fe- 
males, plus the narrower convergent fronts 
could justify a different subgenus than for hir- 
tipalpis and brevirostris when more material be- 
comes available for assessment. 


Species Incertae Sedis 


Mycteromyia robusta Kroeber 
Mycteromyia robusta KROEBER, 1930b:219. 


The description of robusta appears to indicate 
it as belonging to the genus Fidena, NOT Myc- 
teromyia, based on such characters as: Front 
widely divergent below, not parallel-sided or 
convergent as in Mycteromyia; palpi differ in 
shape—blade or knife-shaped; body predomi- 
nantly black, the legs red; abdominal pattern 
unlike Mycteromyia with first three tergites red- 
dish brown on sides, remainder black with pale 
incisures; venter reddish brown with pale hairs. 
The wings are stated only to be fumose without 
mention of “‘punctaugen’’ that are always men- 
tioned by Kroeber in other species of the group. 
Type locality and distributional data were not 
given. 

KEY TO SPECIES 


la. Face produced, snoutlike (Figs. 2A, 3A 
& D;4A & C; 6A & B; 7A & B; 8A,B, 
& 1; 9B, F, & N); proboscis elongated, 
about 2 wing length, labella chitinized; 
apical segment of palpus with short 
hairs, and an elongated dorsal *‘sen- 
sorial’’ groove (Figs. 2C; 3C; 4E; 7C; 
9C, & P); abdomen relatively attenuat- 
ed caudally, female in profile with cerci 
longer than St. VIII (Fig. 3£); in male, 
with genitalia bulbous (Figs. 1D & 
2E) (genitalia distinctive in both sexes 
on tribal level. Tribe Mycteromyiini 
new (Mycteromyia Philippi, Promyc- 
teromyia new genus, Silvestriellus 
Brethes) "1s. 
1b. Face shorter, not snoutlike (Figs. 10A, F, 
K, & M); proboscis extending less than 
Y wing length, labella fleshy; palpus 
more hairy with a short subapical *“‘sen- 
sorial’’ pit (Fig. 10G & L); abdomen 
with normal tabanid appearance, gen- 
italia with normal pangoniine struc- 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


ture, not terminally bulbous in males. 
Tribe Pangoniini (Caenopangonia 


Kerocben) ieee re ee 16 "=Sb: 


2a. Abdomen (female) almost entirely red- 
dish orange with narrow median dark 
line (Fig. 9D); size small (length 9 
mm, wing 8 mm) (Argentina; Buenos 
NITES) ie ee ee PT: oe 
2a S. flaviventris (Barretto and Duret) 

2b. Species with different coloration and pat- 


tern, generally larger than 9 mm long 3 9a. 


3a. Species with distal article of palpus very 
short (0.4-0.6 mm) (Figs. 9FE, J, & O); 


ratio palpus:wing over 1:11 ____-____- 4 
3b. Distal article of palpus over 0.7 mm; ratio 9b. 
palpusswing less than 1:).3 922. 6 


4a. Length 15 mm, wing 11—11.5 mm; color 


reddish brown (southern Argentina) 10a. 


pipe aad S. patagonicus Brethes 
4b. Length between 11-12 mm, wing 8-10.5 
mm; darker brown or dark grayish in 
COLO meee te nie 8 Pie ey eet ee 5 
Sa. Species dark brown, with mid-tergal tri- 
angles well defined, and incisures with 


pale yellow hairs; wing subhyaline, 10b. 


length 10 mm; ratio palpus: wing 1:18— 

24 (Argentina; Chubut and Nequen _ 

ee teoretis ‘aie tat S. schlingeri new species 
5b. Species dark grayish brown, without 

such well-defined triangles; wing veins 


brown margined, length about 10.5 Ila. 


mm; ratio palpus: wing 1:17 (Argentina: 
Buenos Aires)! = ee 
Aaa Sod S. martinezi (Barretto and Duret) 
6a. Species pale yellowish to dark brown; 
large, over 15.5 mm in length —___ 4 
6b. Species light to blackish gray, sometimes 
yellow laterally; size between 10.5—15 
ITD lhl Rel Sa RR ca De pt 9 


7a. Wings strongly fumose, length between I1b. 


13-17 mm; body deep brown; palpi sa- 
LYE CS) re OVE 6 Nae at ale eS ae ee ee ee 8 


7b. Wings very light, body yellowish to ia: 


tawny brown, with dark brown, flat- 
tened spot on each tergite (Fig. 7D), 
palpi not saber-shaped (Fig. 7C) (Chile: 
MAL CA) ig eet a P. xantha new species 
8a. Beard brownish gray. Visible male cerci 
elongated, subtriangulate (Fig. 8F); fe- 


male caudal spermathecal ducts elon- 12b. 


gated (Fig. &8/) with large diverticles 


well sclerotized on the cephalad por- 13a. 


449 


tions (Fig. 8K) (central Chile) —_____ 
Pah esas Sree NO M. conica (Bigot) 
Beard white. Visible male cerci robust, 
subtrapezoidal (Fig. 7G); female cau- 
dal spermathecal ducts short and ro- 
bust, with small unsclerotized diverti- 
cles on caudal portions (Fig. 7/), and 
T. IX short (Fig. 7/) (Chile: Valparaiso 
and Santiago) Sue. Aa 
1 Vem: M. etcheverryae new species 
Species blackish above and below, T. I- 
II with contrasting pale pile (Fig. 1A) 
(Chile: Coquimbo and Atacama) _____- 
AFR EE th ivtyes Bah P. philippii (Philip) 
Species grayish brown, light to dark gray 
or yellow; T. I-II not with special pale 
Pilosity=s. Vee wt Ee Me 10 
Small species, wing length 8.5—9 mm; pal- 
pi ‘short, under” 1-0 mm: ratio 
palpus:wing 1:9-11.5; body gray with 
erect whitish-yellow hairs; visible male 
cerci rounded or truncated (Fig. 6D) 
Chile: ACoquimbo)e i! Whee ss 
REET OSES SIRE _P. penai new species 
Larger species, wing over 9.3 mm (except 
some males of cinerascens); visible 
cerci trapezoidal-shaped (Fig. SA) and 
palpi longer, over 1.0 mm; ornamen- 
tation variable, if gray then with erect 
Dale hairst es: See oe eee 11 
Body coarsely pale pilose dorsally with 
accentuated woolly appearance in 
males overlying light brown integu- 
ment; beard and thoracic hairs whitish; 
female with a mid-abdominal row of 
narrow, pale triangles flanked by 2 sub- 
median stripes of reddish-brown pilos- 
ity (Fig. 4F) (Chile: Valparaiso and 
Santiago): 25) 2 a P. eriodes (Philip) 
Body without this combination of char- 
acters, sometimes yellowish or, if gray, 
not with woolly appearance ________-- 12 
Abdomen mostly gray without yellowish 
sides dorsally and with small subme- 
dian paired spots on T. I-IV; male cer- 
ci wide, and little constricted (Fig. SG); 
female, length about 13 mm, wing 11 
mn (Chile: Aconcagua)’ 22222)". s- 
EROS Ie P. pechumani new species 
Abdomen gray with yellow sides dorsal- 
Loh wee 1 ott Bets Bay 13 
Male with visible cerci rounded (Fig. 6G); 


450 


female, length over 13.5 mm, wing over 
12 mm (Chile: Santiago and Curico) 
Seis aicyuelt umes P. murina (Philippi) 
Male with visible cerci obviously pointed 
(Figs. 2F'; 31; & 5A); female length less 
than 12.5 mm, wing less than 12 mm 


13b. 


14a. Male abdomen predominantly yellow in- 
cluding yellow-brown venter; female 
darker and tergites more gray laterally 
(Chile\Coquimbo) S28 2.288 
P. galbina new species 
14b. Male abdominal tergites not totally yel- 
lowish, at least T. II-VI with promi- 
nent, dark gray basal bands; female, 
likewise but venterdark grayishbrown 15 
Male with very long and pointed visible 
cerci (Fig. 3J & K); female, abdomen 
dark laterally (Chile: Coquimbo) ______ 
Peel ee P. derocerca new species 
15b. Male cerci shorter and less pointed, trap- 
ezoidal-shaped (Fig. 5A); female ab- 
dominal triangles crossing tergites and 
with relatively wide bases, flanked by 
paired submedian grayish-brown dash- 
es, laterally gray with white hairs on 
outer corners (Chile: Santiago and Ata- 
cama) P. cinerascens (Bigot) 
16a. Abdomen light reddish brown with gray 
incisures; legs unicolorous, yellowish 
red (Chile: Osorno and Arauco) _______ 
Pie tS eee C. brevirostris (Philippi) 


Sa. 


Genera 
Promycteromyia, Mycteromyia, and Silvestriellus (Tribe Myc- 
teromyiini). 
1) Thorax, wings, and fore legs elongated (especially the 
coxae). 
2) ‘‘Sensorial area’’ of palpus situated in a frontolateral 
groove (Figs. 2C; 3C; 4E; & 7C). 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


16b. Abdomen dark brown, occasionally with 
light-yellow or gray isolated spots; legs 
with at least femora darker than pre- 
Ceding, 2¢.c.00t.. Lalmenal) Sane 1 
17a. Abdomen dark brown with yellow pos- 
terolateral areas about % width of ter- 
gites in both sexes; face moderately 
produced (Fig. 10K & M); male disti- 
styli pointed (Fig. 10N) (Chile: Coquim- 
DO) pret: teh Pati C. asper (Philip) 
17b. Abdomen very dark brown, light outer 
corners of tergites usually more re- 
duced, at least in males; face in both 
sexes less produced (Fig. 10A); male 
dististyli truncated (Fig. 10D & E) 
(southern Chile and Argentina) 
C. hirtipalpis (Bigot) 


DISCUSSION AND CONCLUSIONS 


Based on exosomatic characters and espe- 
cially on the morphology of the genitalia, we 
consider that the species previously assigned to 
Mycteromyia constitute a polyphyletic group. 

In the following outline, the differences be- 
tween these species are presented. They were 
formerly grouped because of the presence of 
hinged proboscides, combined with other char- 
acters that appear disparately in other Tabani- 
dae, such as dichoptic eyes in males, presence 
in wings of isolated clouds, and of the first pos- 
terior (R;) cells closed, and the nonhaematopha- 
gous habits of females. 


Genus 
Caenopangonia (and genera of other tribes described in Pan- 
goniinae). 
Thorax, wings, and fore legs not unusually elongated. 


‘“‘Sensorial area’’ of palpus reduced to subapical pit (Fig. 
10G). 


Males 


3 


— 


Epandrium bulbous, exposed beyond segment VII (Figs. 
1A & 2E), basally moderately concave (Figs. 1C; 2H; 3K; 
SB & H; & 6E & H) or straight (7J; 8H; & 9/). 

Cerci projected basally into inner epandrium (Figs. 1C; 
2H; 3K; 5H; 6E & H; 7J; 8H; & 91); external portions 
greatly variable in shapes (Figs. 1B; 2F; 3/; SA & G; 6D, 
G,I, & 8F). 

Basistyle inwardly membranous (Figs. 1D; 21; 6F; 7E; 8G; 
& 9H), and with conspicuous styli (also sometimes with 
tuberosities) (Figs. 1D; 2/; 3B; 4F; & 7E & F). 
Gonapophyses ventrobasally extended cephalad (Figs. 
1D; 8G; & 9H). 

Dististyli subtriangulate, acute apically and concave dor- 
sally, frequently with emarginations (Figs. 1D; 2J & J; 3L 
& M; 4B; 5C & 1; 6F & 1; 7E & F; 8G; & 9H). 


4 


~— 


5 


~— 


6 


— 


3 


~— 


Epandrium normally enclosed within segment VII with deep 
concavity (Fig. 10C). 


Cerci not so projected (Fig. 10C); external portions very con- 
stant in shapes. 


Basistyle inwardly not membranous (Figs. 10D & N), and with 
neither styli nor tuberosities. 


Gonapophyses ventrobasally not so extended (Figs. 10D & 
N). 
Dististyli subcylindrical (Figs. 10D, E, & N). 


COSCARON & PHILIP: REVISION OF MYCTEROMYIINI 


451 


Females 


8) Genital fork lightly sclerotized basally (Figs. 3G; 5D; & 
7K), caudal spermathecal ducts elongated (Figs. 3G; 5D; 
& 8J) or thickened (Fig. 9K), generally with peculiar scle- 
rotized portions cephalad (Figs. 1E; 3G; 4G; 5D; & 8K). 
9) St. VIII with lateral emarginations on base (Figs. 4H & 
SE). 
10) T. IX frequently with large lateral expansions (Figs. 1F; 
3A; 4I; SF; GJ; TL; & 8L). 
11) T. X frequently undivided (Figs. 1F'; 3H; 4/; 5F; & 6/J/). 


We consider that this combination of tabulat- 
ed characters justifies the placement of the gen- 
era Promycteromyia, Mycteromyia, and Silves- 
triellus in a separate tribe. We had considered 
establishing a new subfamily, based especially 
on the distinctive morphology of the genitalia, 
but our ignorance of the immature stages, and 
the fact that we have not accomplished an an- 
ticipated detailed worldwide revision of the 
tribes have restrained us from this major action 
for the present. 

Through the above analyses, we conclude that 
some of the characters, like the shapes of the 
dististyli, basistyle membranous inwardly, the 
presence of styli on the basistyle (plus its ceph- 
alad projection on ventral surface) are charac- 
ters shared in common with the Pelecorhynchi- 
dae, and this reaffirms close relationship with 
primitive Tabanidae. Some characters that are 
only present in the new tribe Mycteromyiini, 
and which reinforce our hypothesis that this is 
a monophyletic group, are the shapes of the con- 
spicuous caudal spermathecal ducts, the baso- 
lateral emarginations of St. VIII, and the shapes 
of the dististyli. 

Minor but quite evident characters, such as 
the presence or absence of tuberosities, the di- 
vided or undivided T. X, the shapes and struc- 
tures of caudal spermathecal ducts, the shapes 
of dististyli, aedeagus, and cerci, and the shapes 
and sizes of maxillary palpi, made possible this 
revision of the supraspecific taxa. 


LITERATURE CITED 


BARRETTO, M. P., AND J. P. DureT. 1954. Sobre alguns 
tabanidas argentinos, com a descricao de tres novas espé- 
cies de Mycteromyia Phil. (Diptera, Tabanidae). Rev. Bra- 
sil. Entomol. 1:203-212. 

Bicor, J. 1858. Dipteres nouveaux provenant du Chili. Ann. 
Soc. Entomol. Fr. (1857)(3)5:277-308. 

. 1892. Descriptions de Dipteres nouveaux. Mém. Soc. 
Zool. Fr. 5:602-691. 

BRETHES, J. 1910. Dipteros nuevos 6 poco conocidos de Sud- 
America. An. Mus. Nac. Buenos Aires 20 (=Ser. 3, 
13):469-484. 


Genital fork well sclerotized basally with normal caudal sper- 
mathecal ducts, without special sclerotized portions (Fig. 
10/). 


St. VILL without lateral emarginations (Fig. 1/). 
T. IX without such expansions (Fig. 10/). 


T. X always divided (Fig. 10/). 


. 1921. Los Tabanidos del Plata. Estudios 21: 1-79. 

ENDERLEIN, G. 1922. Ein neues Tabanidensystem. Mitt. 
Zool. Mus. Berl. 10:333-351. 

FAIRCHILD, G. B. 1971. Family Tabanidae. Jn, A catalogue 
of the Diptera of the Americas south of the United States. 
Mus. Zool., Univ. Sao Paulo, Brasil, fasc. 28. 163 pp. 

Hack, W. H. 1953. Segunda contribucion al estudio de los 
tabanidos argentinos (Diptera). An. Inst. Med. Reg. 3:339 
347. 

KRogEBER, O. 1930a. Tabanidae. Diptera of Patagonia and 
South Chile 5:106—161. 

. 1930b. Die Tribus Pangoniini der neotropischen Re- 

gion. Zool. Anz. 89:211-228. 

1934. Catalogo dos Tabanidae da America do Sul e 
Central, incluindo o Mexico e as Antilhas. Rev. Entomol. 
(Rio de Janeiro), 4:222-276. 

MACKERRAS, I. M. 1955. The classification and distribution 
of Tabanidae (Diptera). II. History: Morphology: Classifi- 
cation: Subfamily Pangoniinae. Aust. J. Zool. 3:439-S11. 

Puitip, C. B. 1958. Descriptions of new neotropical Taban- 
idae mostly in the California Academy of Sciences (Dip- 
tera). Pan-Pac. Entomol. 34:63-76. 

1968. The types of Chilean species of Tabanidae 

(Diptera) described by Dr. R. A. Philippi. Rev. Chil. En- 

tomol. 6:7-16. 

1977. Designation of Pangonia conica Bigot, 1857, 
not Philippi: 1865, as type-species of Mycteromyia Philippi, 
1865 (Insecta, Diptera, Tabanidae). Z.N.(S.) 2199. Bull. 
Zool. Nomencl. 34:187-188. 

PHILippI, R. A. 1865. Aufzahlung der chilenischen Dipteren. 
Verh. Kaiserl.-KGnigl. Zool.-Bot. Ges. Wien 15, Abh.:595— 
782. 


RESUMEN: “Revision de las especies incluidas 
en Mycteromyia por diferentes autores (Diptera- 
Tabanidae).”’ 


Estudios morfologicos, especialmente basa- 
dos en la genitalia, evidencian que las especies 
de tabanos primitivos del sur de América del Sur 
incluidos en Mycteromyia Philippi 1865, consti- 
tuyen un grupo parafilético. 

En base a ello se han redistribuido taxono- 
micamente de la siguiente manera: De las 13 es- 
pecies asignadas anteriormente al género Myc- 
teromyia ubicado en la primitiva tribu Scionini 
(subfamilia Pangoniinae), 3 son retenidas con 
reservas en dicha tribu en el revalidado género 
Caenopangonia Kroeber, nominadas hirtipalpis 


452 


(Bigot) (especie tipo), brevirostris (Philippi), y 
asper (Philip). Dos especies, Pangonia obscu- 
ripennis Philippi y M. robusta Kroeber no se 
han visto. Por las descripciones consideramos 
que no pertenecen a este grupo y por ello no se 
incluyen en esta revision. 

Las restantes 8 especies, ademas de otras 7 
anteriormente no descriptas, son aqui asignadas 
a la nueva tribu Mycteromyiini de la subfamilia 
Pangoniinae con 3 géneros: Promycteromyia 
n.gen. para Promycteromyia philippii (Philip) 
(especie tipo, nueva combinacioén), P. galbina 
n.sp. (macho y hembra, Chile), P. derocerca 
n.sp. (macho y hembra, Chile), P. eriodes (Phil- 
ip) (nueva combinacion), P. cinerascens (Bigot) 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 19 


(nueva combinacion), P. pechumani n.sp. 
(macho y hembra, Chile), P. penai n.sp. (macho 
y hembra, Chile), P. murina (Philippi) (nueva 
combinacion), y P. xantha n.sp. (macho, Chile); 
Mycteromyia para Mycteromyia conica (Bigot) 
(especie tipo) y M. etcheverryae n.sp. (macho 
y hembra, Chile); y Silvestriellus para Silves- 
triellus patagonicus Brethes (especie tipo), S. 
schlingeri n.sp. (macho y hembra, Argentina), 
S. martinezi (Barretto y Duret) (nueva combi- 
nacion), y S. flaviventris (Barretto y Duret) 
(nueva combinacion). Mycteromyia bejaranoi 
Barretto y Duret es sinonimizada con D. hirti- 
palpis Bigot. 


iP ‘en | 
i a - ao 
7 i 
Baa at ogame 
aime ibiiacees & 


——o- 4 


= _ 


” hae 7 
tly" ners © = oer > 7 


7 


-_, 
« 

4 my) 

ys ag 

7 as 


‘pee 1), FY. MRT Fee 
SOGPE, CMPe))*) Theipeeling 
ee Pus cagi s¢ We ed 

1 Astvyt bie Sreee sf Pee Ib ae tis 

~ us (have -Sae jrpltty Lr aus") 

a ee > AINA -> an ieee 


a a 4 a : ve @ asl eG “Goa. ah; oe 


i as 4 a e.Peaee “7? Qe © Ve Pee 


, ey Aneta pecehia ae eae | Oa) nell Ppa 
eek ate, Meek asiny «yp 


i 
NH 


PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 
FOURTH SERIES 


Vol. XLI, No. 20, pp. 453-473; 9 figs., 3 tables. 


January 24, 1979 


THE SCORPIONFISH GENUS MINOUS (SCORPAENIDAE, MINOINAE) 
INCLUDING A NEW SPECIES FROM THE INDIAN OCEAN 


William N. Eschmeyer and Leon E. Hallacher' 
California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 


and 


Kaza V. Rama-Rao 
Marine Biological Station, Zoological Survey of India, 69 Santhome High Road, Madras 600 028, India 


ABSTRACT: 


The scorpionfish genus Minous, the only genus in the subfamily Minoinae, is reviewed. The 


nominal genera Corythobatus Cantor, Decterias Jordan and Starks, Lysodermus Smith and Pope, and Par- 
aminous Fowler are treated as synonyms of Minous Cuvier. Nine species are recognized, of which M. 
dempsterae is described as a new species occurring from the Gulf of Oman to western India. The genus is 
known from the western Pacific and the Indian Ocean and the species are found on soft bottoms in depths 
from about 10 to 420 m. Descriptions, synonymies, figures, and ranges are given for each species, and a key 


to the species is provided. 


INTRODUCTION 


Scorpionfishes of the genus Minous occur 
only in the Indo-West Pacific faunal region, from 
Japan to Australia westward to the western In- 
dian Ocean and Red Sea. Species are mostly 
coastal in distribution and live at depths between 
about 10 and 420 meters. Most, if not all, live on 
mud or sand bottom in marine waters and appear 
to be common in these habitats. They are small, 
scaleless fishes, rarely exceeding 15 cm in total 
length; some grow only to about 7.5 cm. Mem- 
bers of the genus are characterized by having 
the lowermost ray of the pectoral fin separated 
from the rest of the fin; these free rays are cov- 
ered distally with a peculiar “‘cap’’ (or “‘glove’’), 


' Present address: 198 Park Ave., Monterey, California. 


and are evidently used for ‘‘walking”’ on the bot- 
tom. 

Our knowledge of the species of Minous is 
based on regional studies, and no previous au- 
thor has attempted to treat all species. Little is 
known about these fishes, and their higher-cat- 
egory placement among other scorpionfishes is 
uncertain. They are frequently placed in the 
stonefish family, Synanceiidae (e.g., as by Herre 
1951; Smith 1957; and de Beaufort 1962), but 
Jordan and Starks (1904), Fowler (1938a), and 
Matsubara (1943) among others felt that they 
constituted their own subfamily Minoinae of the 
Scorpaenidae. We distinguish the subfamily Mi- 
noinae from the Synancelinae and Choridacty- 
linae (nimicus and Choridactylus) by the lack 
of accessory lateralis pores on the body in the 
Minoinae, among other features (see Eschmeyer 
and Rama-Rao 1973). 


[453] 


454 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


Some spécies of the genus Minous are inter- 
esting because of their presumed commensal re- 
lationship with gymnoblastic hydroids. Alcock 
(1892) recorded a case of commensalism be- 
tween Stylactis minoi Alcock and Minous iner- 
mis Alcock from India. Franz and Stechow 
(1908) and Franz (1910) reported Stylactis minoi 
on Minous spp. from Japan. Also, at least some 
species are known to have venomous fin spines. 


ACKNOWLEDGMENTS 


Most of the specimens used in the study were 
provided by the Smithsonian Oceanographic 
Sorting Center or were ones available to us in 
the collections of the California Academy of Sci- 
ences (CAS) and the Zoological Survey of India 
(ZSI). Loans were made by Dr. T. Abe, Uni- 
versity of Tokyo (UT); the United States Na- 
tional Museum (USNM); Dr. G. R. Allen, West- 
ern Australian Museum, Perth (WAM); Dr. R. 
M. Bailey, University of Michigan Museum of 
Zoology (UMMZ); Dr. J. E. Bohlke, Academy 
of Natural Sciences of Philadelphia (ANSP); Dr. 
R. K. Johnson, Field Museum of Natural His- 
tory (FMNH); and Dr. A. D. Welander, Uni- 
versity of Washington, College of Fisheries 
(UWCF). A visiting-scientist fellowship provid- 
ed by the California Academy of Sciences and 
a grant from the Smithsonian Institution through 
the U.S. Foreign Currency Program allowed 
Rama-Rao to participate in this study. We are 
grateful to the Zoological Survey of India for 
permitting him to travel to the United States. A 
trip to overseas museums by Eschmeyer was 
supported by National Science Foundation grant 
GB-15811. We thank the following persons for 
assistance to Eschmeyer during visits to their 
museums: Drs. M. L. Bauchot and J. C. Hureau, 
Museum National d’Histoire Naturelle, Paris 
(MNHN); Dr. M. Boeseman, Rijksmuseum van 
Natuurlijke Historie, Leiden (RMNH); Drs. C. 
Karrer and K. Deckert, Zoologische Museum, 
Humboldt University, Berlin (ZMHU); Dr. A. 
G. K. Menon, Zoological Survey of India, Cal- 
cutta (ZSI); Dr. J. Paxton, Australian Museum, 
Sydney (AMS); and A. Wheeler and staff, Brit- 
ish Museum (Natural History), London 
(BMHN). Lillian J. Dempster helped with lit- 
erature and translations, and she provided com- 
ments on the manuscript. Drawings were made 
by Katherine Smith, and photographic assis- 
tance was provided by Kenneth E. Lucas, 
Maurice Giles, and Lloyd Ulberg. Franz B. Stei- 


ner, CAS Field Associate, made a special effort 
to collect specimens from India, Thailand, Tai- 
wan, and the Philippines. Others at CAS assist- 
ing in the study were M. Barbour, W. I. Follett, 
J. Gordon, T. Iwamoto, B. Meinhard, S. Poss, 
B. Powell, C. Ruark, P. Sonoda, and B. Wese- 
mann. 


METHODS 


Methods are those used by Eschmeyer (1969) 
and differ little from those used to describe te- 
leost fishes. In species of the genus Minous, the 
last element of the dorsal and anal fins is single, 
separate, and well spaced from the preceding 
ray, and it is counted as a full ray; all soft rays 
of the dorsal and anal fins are unbranched. Pec- 
toral fin length is measured from the base of the 
uppermost ray to the tip of the longest ray. 


SUBFAMILY MINOINAE 


Minoinae JORDAN AND STARKS, 1904:93 (in key; for Minous 
and Decterias). FOWLER 1938a:51 (in key). MATSUBARA 
1943:398-399 (description; internal features). 


REMARKS.—Only the genus Minous, with its 
synonyms Corythobatus, Decterias, Lysoder- 
mus, and Paraminous, belongs to this subfami- 
ly. The subfamily is diagnosed under the genus 
below. 


Genus Minous Cuvier 


Minous CUVIER in CUVIER AND VALENCIENNES, 1829:420 
(type-species Scorpaena monodactyla Bloch and Schnei- 
der, 1801, by subsequent designation of Bleeker 1876). 

Corythobatus CANTOR, 1849:1027 (replacement name for Mi- 
nois Hubner, 1816, Lepidoptera). 

Decterias JORDAN AND STARKS, 1904:154 (type-species Min- 
ous pusillus Temminck and Schlegel, 1843, by original des- 
ignation). 

Lysodermus SMITH AND Pope, 1906:483 (type-species Lyso- 
dermus satsumae Smith and Pope, by original designation). 

Paraminous FOWLER, 1943:68 (type-species Paraminous 
quincarinatus Fowler, by original designation). 


NOMENCLATURAL REMARKS.—Reasons for 
including the nominal genera listed above as 
synonyms of Minous are as follows. The genus 
Corythobatus was proposed as a substitute 
name for Minous, which Cantor thought was 
preoccupied by Minois Hiibner, but according 
to Article 56(a) of the International Code of Zo- 
ological Nomenclature, Minous and Minois are 
both available generic names as they differ in 
spelling. Lysodermus was based on an abnormal 
specimen of M. monodactylus, a specimen with 
7 instead of the normal 9-11 dorsal spines. Dec- 


ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 455 


terias and Paraminous were established on 
characters, such as flexible dorsal spines and 
short head spines, which do not warrant sepa- 
rate genera. Species may be sorted into groups 
based on different characters, but no sharp di- 
vision is noted between groups of species. Min- 
ous monodactylus, M. versicolor, and M. quin- 
carinatus have the first dorsal spine long and 
well separated from the second spine, while the 
other species have a small first dorsal spine 
which is situated very close to the base of the 
second spine; the long first dorsal spine in these 
three species results from a loss of the first small 
spine as evidenced by the anterior vertebrae, 
neural arches, and pterygiophores. Minous pus- 
illus is the only species with hairlike dorsal 
spines; M. inermis has thin and flexible dorsal 
spines, while the others have sharp, firm spines, 
but in some species the spines are thin and flex- 
ible in the young (e.g., M. trachycephalus). Par- 
aminous was established for two species on the 
basis of their having short head spines, but this 
condition also occurs in other species. On the 
basis of head spination, two groups, one with 
five species and one with four, are evident: those 
with long head spines (especially the preoper- 
cular and posterior lachrymal spines) and those 
with short spines. Five species have a gas blad- 
der and four apparently do not, but we can not 
correlate this with other morphological differ- 
ences. We therefore conclude that all species 
belong in the genus Minous. 
DraGNosis.—Dorsal spines 8-12, dorsal soft 
rays 8-14; total dorsal fin elements 19-24. Anal 
fin with 2 spines and 7-11 soft rays, total 9-13; 
spines almost indistinguishable from soft rays. 
Pectoral fin normally with 12 rays in all species; 
lower pectoral ray completely free from rest of 
fin, slightly enlarged, and fitted at its tip with a 
cuticular “‘cap’’ (this cap frequently rubbed off 
in preserved specimens). Pelvic fin with | spine 
and 5 soft rays. Soft rays of all fins unbranched. 
Second suborbital bone (= third circumorbital 
bone) becoming wider posteriorly and firmly at- 
tached to preopercular bone. Scales absent, ex- 
cept for tubes of lateral line. No accessory lat- 
eralis pores on body. Vertebrae 24-27. Gas 
bladder present or absent. First 2 neural spines 
point forward; first 2 or 3 dorsal fin pterygio- 
phores (2 if first dorsal spines is lost) inter- 
spersed between second and third neural spines. 
Hypurals coalesced into 2 plates. Teeth in jaws 
small; palatine teeth absent; vomerine teeth 


present. Upper part of eyeball and lower jaw 
with dermal cirri; no tentacles or skin flaps on 
body. 

SPECIES CHARACTERS.—The nine species of 
the genus may be separated with various com- 
binations of characters. Several species can be 
identified merely by the coloration on the inside 
of the pectoral fin. As in many other scorpion- 
fishes, display of bright colors in the pectoral fin 
axil as a fright response probably occurs in spe- 
cies of Minous. The first two dorsal spines are 
either about the same length and normally 
spaced or the first is very short and close to the 
base of the second spine. The preopercular 
spines are either well developed or moderately 
developed, usually six in number, with the long- 
est (second from top and immediately posterior 
to suborbital ridge) reaching to the end of the 
opercle or reaching only about halfway across 
the opercle. The posterior lachrymal spine lying 
over the maxillary is very long and bayonet- 
shaped in some species or it is short and stubby 
or intermediate in other species. Body color- 
ation was found to be useful in separating some 
species. One species has the dorsal spines nearly 
hairlike, while they are flexible or stout in the 
others. One species has an especially long pec- 
toral fin. Meristic features, particularly of dorsal 
and anal fin rays and vertebrae, show some dif- 
ferences among the species. 

DISTRIBUTION.—Species of the genus Minous 
are restricted to the Indian Ocean and the west- 
ern Pacific as presented in Figure 1. We suspect 
that further trawling will reveal that some spe- 
cies have wider ranges than now known. They 
can be classified as coastal species, occurring on 
mud or sand bottom. 


Key to the Species of the Genus Minous 


la. First dorsal spine equal to or longer than 
second dorsal spine, well separated 
from the base of the second spine ___- 


nN 


lb. First dorsal spine shorter than second 
dorsal spine, usually less than half the 
second and sometimes extremely small, 
close to second spine at base___________- 4 


2a. Caudal fin without transverse dark bars; 
inside of pectoral fin with dark brown 
stripes overlying the rays (Fig. 2c); pos- 
terior lachrymal spine moderate, about 
twice length of anterior spine and not 
bayonet-shaped (Fig-:2a)) 2 2 
ih rR M. quincarinatus (Fig. 3a) 


456 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


20° 40° 60° 80° 100° 120° 140° 160° 


a PHILIPPINE 
Is. 


hea 


NICOBAR, 
16. 


re CEYLON 
MALDIVE™: = 
1s 


a 


SEYCHELLES 1S. 


CHAGOS |” 
ists 


- coccineus 
- dempsterae 
- inermis 


+ monodactylus 


il ° 
20° 40° 60° 80° 100° 120° 140° 60 


Ew 


- pictus 


- pusillus 


+ quincarinatus 
- trachycephalus 
- versicolor 


Ficure 1. Distribution of the species of the genus Minous. 


ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 457 


DLO 


FIGURE 2. Diagram of lachrymal spines (in black) in (a) Minous quincarinatus and (b) M. monodactylus; coloration on 
inside of pectoral fin in (c) M. quincarinatus, (d) M. coccineus, and (e) M. dempsterae. 


2b. Caudal fin with transverse dark bars; in- 4b. Dorsal spines strong or moderately 
side of pectoral fin mostly plain col- strong, or thin and flexible, not hairlike 
Gledeenoteassinahicure 2C4sDOStehOl may) | 2a. 2 Bane ee ee 5 
lachrymal spine long and bayonet- Sa. Pectoral fin long (more than 50% SL), 
Shapedi(iign.2b)! 2... 2 ee 3 reaching to over end of anal fin________- 
3a. Dorsal spines usually 9; caudal finusually 4 =e __M. inermis (Fig. 5) 
with 3 or 4 transverse wavy bands; soft 5b. Pectoral fin moderate (less than 46% SL), 
dorsal fin with wavy bands_______________- reaching to middle of anal fin as a max- 
3) see eee ee M. versicolor (Fig. 3b) LULU Test ee ee ie ee eee ee. 6 
3b. Dorsal spines usually 10 or more; caudal 6a. Soft dorsal fin with 8-10 rays, usually 10; 
fin with 2 broad vertical dark bars; soft anal fin spines plus soft rays total 9-11, 
dorsal fin intensely black on anterior usually 10; lachrymal spines about 
nalipie: a. M. monodactylus (Fig. 3c) same length; caudal fin rays usually 
4a. Dorsal spines weak and hairlike _____-_____- with alternating dark and pale color- 


RE eee eee eee M. pusillus (Fig. 4) ation __________M. trachycephalus (Fig. 6) 


458 


TABLE 1. 


Species 


. quincarinatus 

. versicolor 

. monodactylus' 
. pusillus 

. Inermis 

. trachycephalus 
. coccineus 
pictus 

. dempsterae 


= = << S 


ae Tie | 
— Me n— 


' One abnormal specimen with VII spines. 


6b. Soft dorsal fin with 11-13 rays, usually 
12; anal fin spines plus soft rays 11-13, 
usually 12; posterior lachrymal spine 
much longer than anterior lachrymal 
spine; caudal fin pale, with no dark 
markings 
Inside of pectoral fin with irregular black 
spots on a pale background (Fig. 2d) _ 
M. coccineus (Fig. 7) 
Inside of pectoral fin without irregular 
black spots 
Inside of pectoral fin with stripes radiating 
distally along course of fin rays (similar 
to M. quincarinatus, Fig. 2c) 
ee. I ees. M. pictus (Fig. 8a) 
Inside of pectoral fin with pale spots on 
a dark background (Fig. 2e) 
M. dempsterae (Figs. 8b, 9) 


ae 


7b: : 


8a. 


8b. 


Minous quincarinatus (Fowler) 
(Figures 1, 2a,c, 3a; Tables 1-2) 


Paraminous quincarinatus FOWLER, 1943:69-70, fig. 13 (orig- 
inal description; type locality southern Japan, 30°12'N, 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


DorSAL FIN RAY COUNTS IN SPECIES OF GENUS Minous. 


Total rays 
il il 14 197 920), 9217 22552324 
— Le: 2 —_-_ — De 12 1 — 
ay lil 5 — — Ay 512 3 — — 
30 2 — ~ 1 6 36 —- — — 
2—-— — —_ — 8 7 -—- — — 
— 2 8 9 —- — — A a2 3 
—- — — — 2, 6 6 — — — 
4 18 — — —_- — 1 4 17 — 
e238 2 — —- —- — DY 628) 1 
3 — —- —- — 1 25 2 


130°44'E, 84 fms [154 m], ALBATRoss dredge sta. 4930, 15 
Aug. 1906; holotype USNM 99515). 

?Minous inermis: FRANZ AND STECHOW 1908:753 (Sagami 
Bay, Japan; infested with hydroid Stylactis minoi Alcock). 

?Minous adamsi var inermis: FRANZ 1910:73, pl. 9, fig. 70 
(brief description; hydroids growing on specimens; 2 from 
Misaki, Japan). 

Minous inermis: JORDAN, TANAKA, AND SNYDER 1913:247 
(listed; Japan). MATSUBARA 1943:401, 406-409, fig. 139 
(misidentified; good description; comparisons; internal fea- 
tures; Kumano-Nada and Nagasaki, Japan). KAMOHARA 
1952:68 (listed; Japan); 1958a:57 (listed; Japan); 1964:75 
(listed; Japan). 


MATERIAL EXAMINED.—Japan: USNM 99515 (75.0 mm 
SL, holotype of M. quincarinatus), for detailed locality see 
synonymy. 

Taiwan: CAS 13879 (8, 97.0-116), CAS 13880 (1, 79.0, 
cleared and stained), and USNM 218456 (2, 102-108), Taiwan 
Strait, about 26°N, 121°E, about 90 m, bottom trawl, F. B. 
Steiner, 16 June 1971. CAS 13882 (3, 102-108), Taiwan Strait, 
about 25°N, 120°E, about 90 m, bottom trawl, F. B. Steiner, 
Apr. 1971. Additional material: CAS 30003 (1), CAS 30308 (6). 


DESCRIPTION (See also generic diagnosis; 
Figs. 2a,c, 3a; Tables 1—2).—Dorsal fin with 8— 
9 (usually 9) spines and 12-14 (usually 13) soft 


TABLE 2. ANAL FIN RAY AND VERTEBRAL COUNTS IN SPECIES OF THE GENUS Minous. 


Total anal spines 


jandirays! een ee Vertebrae 

9 10 11 12 13 24 25 26 27 
M. quincarinatus — 1 2 12 — — — 3 9 
M. versicolor — 1 17 1 —_— -- 1 8 = 
M. monodactylus 2 3 29 4 — — 1 14 — 
M. pusillus —_— 21 4 —_ — — 16 — — 
M. inermis — l 6 10 2 — — 3 16 
M. trachycephalus 1 20 2 — — 1 6 —_ = 
M. coccineus — — 4 18 i — — 7 — 
M. pictus — — 6 19 1 1 2 8 1 
M. dempsterae — — 11 22 1 — 1 18 1 


ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 459 


Figure 3. (a) Minous quincarinatus, CAS 30308, 105 mm SL, Taiwan. (b) M. versicolor, WAM P4397, 84.5 mm SI, Exmouth 
Gulf, Australia. (c) M. monodactylus, BPBM uncat., 88 mm SL, Madras, India (photo by John E. Randall from a fresh 


specimen.) 


460 


rays; total dorsal spines and rays 21-23 (usually 
22); dorsal spines sharp and strong; first dorsal 
spine long, about equal to second and well sep- 
arated from it. Anal fin with 2 spines and 8-10 
(usually 10) soft rays; spines difficult to distin- 
guish from soft rays; total anal fin elements 10- 
12 (usually 12). Vertebrae 26 (3 specimens) or 
27 (9). Gill rakers total 11-16 on outside of first 
arch, 3-4 on upper arch, 8-12 on lower arch. 
Lateral line pores 16-18. Gas bladder absent. 
Pectoral fin reaching to about beginning of anal 
fin. Head with ridges, spines poorly developed. 
Lachrymal bone with 2 sharp but short spines 
over maxillary; first points down and forward, 
second down and slightly to rear; second spine 
slightly longer than first. 

General body shape and preserved coloration 
as in Figure 3a. First 4 or 5 dorsal spines with 
blackish area at distal part of membrane. Body 
coloration variable. Dorsal half of body brown- 
ish or grayish, with blotchy pale spotted areas, 
and irregular pale stripes. Ventral portion of 
body pale, lacking markings. Anal fin black an- 
teriorly, pale posteriorly. Pectoral fin base whit- 
ish, outside of fin mostly black. Inside of pec- 
toral fin as in Figure 2c; axil pale, usually with 
2 or 3 brown spots; dark stripes radiate out along 
pectoral rays; fin darker distally. Pelvic fin base 
pale, fin blackish distally. Caudal fin pale. 

Measurements for 14 specimens (75.0—-116 
mm SL) in percent of standard length: head 37— 
45; orbit 10-12; snout 13-17; interorbital width 
8-10; postorbital 15-20; pectoral fin 35—46. 

DISTRIBUTION.—Minous quincarinatus is an 
Oriental species known from southern Japan and 
Taiwan in depths ranging from 90-154 m. 


Minous versicolor Ogilby 
(Figures 1, 3b; Tables 1-2) 


Minous versicolor OGiLBy, 1910:111—113 (original descrip- 
tion; type locality Australia, Cape Gloucester, in 26 fms). 
McCuLLocu 1915:162-163, pl. 32, (description; compari- 
son with monodactylus and adamsi); 1929:391 (compiled). 
MARSHALL 1965:428—429, fig. 414 on pl. 57 (brief descrip- 
tion; figure from McCulloch 1929). 

Minous monodactylus: McKay 1964:11 (in text; recorded 
from Western Australia; sting). MEES 1964:48 (versicolor 
probably a synonym; Australian specimens). 


MATERIAL EXAMINED.—AII from Australia. Syntypes of 
M. versicolor: AMS E2736 (2, 71.4-74.1), AMS E2738 (1, 75), 
AMS 113603-—13604 (2, 75.7 —76.5), all from 7 miles (11 km) 
NNE off Bowen, Queensland. 

Gulf of Carpentaria: WAM P5711 (1, 57), Albatross Bay, E. 
Gamberg, 20 Jan. 1962. WAM P5712 (1, 54.3), Albatross Bay, 
E. Gamberg, 22 Jan. 1962. WAM P10827 (1, 55), sta. 1530, R. 
J. McKay, Oct. 1964. 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


Western Australia: WAM P4397 (1, 84.5), Exmouth Gulf, 
R. J. McKay, 19 Oct. 1958. WAM P5714 (1, 79.4), Shark Bay 
area, W. and W. Poole, June—July 1971. WAM P5735 (1. 47), 
Dampier Archipelago, about 6-7 miles (ca. 10-11 km) N of 
Delambre I., B. R. Wilson, 9 June 1960. WAM P7073-75 (3, 
77.1-83.9), Shark Bay, R. J. McKay, 1962. WAM P13934-36 
(3, 52-80), Shark Bay, W. and W. Poole. WAM P14196—97 
(2, 45.8-61.5), Shark Bay, R. J. McKay, June 1962. 


DESCRIPTION (See also generic diagnosis; Fig. 
3b; Tables 1—2).—Dorsal fin with 8—9 (usually 9) 
spines and 11-13 (usually 12) soft rays; total dor- 
sal spines and rays 20-22 (usually 21); dorsal 
spines sharp and strong; first dorsal spine long, 
about equal to second in length and well sepa- 
rated from it. Anal fin with 2 spines and 8-10 
(usually 9) soft rays; spines difficult to distin- 
guish from soft rays; total anal fin elements 10- 
12 (usually 11). Vertebrae 25 (1 specimen) or 26 
(8). Gill rakers total 10-14 on first gill arch, 2-4 
on upper arch, 7-11 on lower arch. Lateral line 
pores 17-19. Gas bladder absent. Pectoral fin 
reaching to middle of anal fin as a maximum. 
Head spines well developed. Lachrymal bone 
with 2 sharp spines over maxillary; first long, 
about half length of second; second very long 
and curved to rear. 

General body shape and preserved coloration 
in Figure 3b. Lower half of body whitish, upper 
half with stripes and oblique bars. Irregular 
stripes extend along lateral line, above and be- 
low. More or less regular oblique bars cross the 
dorsal fin and usually extend onto body. Dorsal 
fin membranes blackish distally. Outer surface 
of pectoral fin dark distally; inner surface of pec- 
toral fin mostly dark gray or black with one or 
two paler areas, without spots or stripes. Pelvic 
fin and anal fin dusky, black distally. Caudal fin 
usually with 4 cross bars. 

Measurements for 19 specimens (45.0—84.5 
mm SL) in percent standard length: head 37-46; 
orbit 11—16; snout 9-12; interorbital width 7-12; 
postorbital 13-21; pectoral fin 35—41; first dorsal 
spine 16-21; second dorsal spine 12-17. 

DISTRIBUTION.—Minous versicolor is the 
only Australian species of the genus and occurs 
from Shark Bay, Western Australia, across 
northern Australia and south on the east coast 
to Queensland. Few depths of capture are 
known and are between 12-64 m. 


Minous monodactylus (Bloch and Schneider) 
(Figures 1, 2b, 3c; Tables 1-2) 


Scorpaena monodactyla BLOCH AND SCHNEIDER, 1801:194 
(original description; no type locality). 


ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 461 


Apistus minous CUVIER, 1829:167—168 (listed; based on Rus- 
sell 1803, pl. 159A). 

Apistus monodactylus CUVIER in CUVIER AND VALEN- 
CIENNES, 1829: fig. 2 on pl. 95 (figure). 

Minous woora CUVIER in CUVIER AND VALENCIENNES, 
1829:42 1-424 (original description; based in part on Russell 
1803:44, pl. 159A, as ‘Woora-minoo’ from Vizagapatam, 
India, and on specimens from Isle-de-France [Mauritius]). 
RICHARDSON, 1846:213 (specimens from China). BLEEKER 
1852a:251—252 (description; Sumatra). 

Scorpaena biaculeata: CUVIER in CUVIER AND VALEN- 
CIENNES 1829:424 (nomen nudum [unpublished name from 
Kuhl and Van Hasselt]; as synonym of M. monodactylus). 

Minous monodactylus; CUVIER in CUVIER AND VALEN- 
CIENNES 1829: 424-426 (description based on specimen 
from Batavia; figured on pl. 95 under name of Apistus mon- 
odactylus). BLEEKER 1849:9 (listed; Java); 1851:19 (listed; 
Indian Archipelago); 1852b:720 (listed; Banka); 1854a:57 
(listed; Sumatra). GUNTHER 1860: 148-149 (synonymy; 
British Museum specimens from India, China, and Borneo; 
M. adamsi as a synonym). BLEEKER 1861:31, 49 (listed; 
Singapore); 1874:87 (listed; Mauritius); 1876b:6, 10-12, 61, 
63-65 (synonymy; description; specimens from Sumatra, 
Pinang, Singapore, Bangkok, Java, Borneo, and Celebes). 
Day 1875:159, fig. 7 on pl. XXXVIII (synonymy; descrip- 
tion; range India to China). BLEEKER 1879a:fig. 2 on pl. 
CCCCXII (color plate); 1879b:16 (listed; Mauritius). DAY 
1889:70-71, fig. 29 (compiled from Day 1875). REGAN 
1905a:20 (M. adamsi and M. echigonius as synonyms; Ja- 
pan); 1908:237 (listed; Maldives in 30 fathoms [55 m]). Jor- 
DAN AND THOMPSON 1914:276—277, fig. 47 (listed; Japan; 
figure from Jordan and Starks, 1904). JoRDAN AND HUBBS 
1925:275 (M. adamsi, M. echigonius, and M. satsumae as 
synonyms; Japan). MACHAN 1930:437 (brief description; 
Padang). KAMOHARA 1930:49 (listed; Japan; common 
name). TANAKA 1931:36 (synonymy; Japan). HERRE 
1932:437 (3 specimens from China). WANG 1935:471-473, 
fig. 46 (synonymy; good description; a specimen from- 
Chefoo; M. adamsi, M. echigonius, and M. satsmae (sic) 
in synonymy). TORTONESE 1937:53 (description; 1 specimen 
from Massaua, Red Sea). HERRE AND Myers 1937:34 (2 
specimens from Singapore). FOWLER 1938b:36 (brief de- 
scription; 1 specimen from Hong Kong). MATSUBARA 
1943:401-404, fig. 138 (synonymy; good description; varia- 
tion; internal features; specimens from Japan). BLEGVAD 
1944:191 (description; 3 specimens from Iranian Gulf; figure 
from Day 1875; distribution). KAMOHARA 1952:68 (com- 
piled; Japan). MUNRO 1955:251, fig. 728 (synonymy; brief 
description; Gulf of Mannar). KAMOHARA 1958a:56 (com- 
piled; Japan); KAMOHARA 1958b: 9 (listed; Japan). DE 
BEAUFORT in WEBER AND DE BEAUFORT 1962:111-113 
(synonymy; description; compiled range). KAMOHARA 
1964:74-75 (compiled). 

Apistes monodactylus: SWAINSON 
Apistes; Minous in synonymy). 
Apistes Russellii: SWAINSON 1839:265 (replacement name 

based on woora-minoo A of Russell 1803). 

Minous adamsii RICHARDSON, 1848:7-8, figs. 4-5 on pl. II 
(original description; type locality Sea of China). 

Corythobatus woora: CANTOR 1849:45 (as type-species of 
Corythobatus). 

Minous Blochi: KAupP 1858:333 (replacement name for Scor- 
paena monodactyla Bloch and Schneider). 

Corythobatus monodactylus: BLEEKER 1865:286 [as 282 mis- 
print] (listed). 


1839:265 (included in 


Minous oxycephalus BLEEKER 1876b:65—66, fig. 2 on pl. 1 
(original description; type locality Sumatra and Amboina). 
Minous oxyrhynchus BLEEKER 1879:fig. 4 on pl. CCCCXVII 
(presumably an incorrect subsequent spelling of Oxycepha- 

lus Bleeker, 1876). 

Minous echigonius JORDAN AND STARKS 1904:151, 153-154, 
174, fig. 14 (original description; type locality Niigata, Ja- 
pan; holotype SU 7380). 

Lysodermus satsumae SMITH AND Pope, 1906:483—485, fig. 7 
(original description; type locality Kagoshima, Japan; ho- 
lotype USNM 55615). JoRDAN, TANAKA, AND SNYDER 
1913:246, fig. 181 (compiled from Smith and Pope 1906). 

Minous adamsi: JORDAN AND STARKS 1904: 151-153 (descrip- 
tion; specimens from Japan). SNYDER 1912:428 (specimens 
from Japan). JORDAN, TANAKA, AND SNYDER 1913:247 (M. 
echigonius in synonymy; range). 

MATERIAL EXAMINED.—Pakistan: CAS 33895 (2, 46.0-— 
53.0), 12 miles (ca. 19 km) from Karachi, 32 m, bottom trawl, 
sandy bottom, F. B. Steiner, 19 June 1975. 

India: CAS 13912 (4, 49.8-59.2), NW India, 21°23'N, 
69°46'E, 18 m, trawl, soft green clay and mud bottom, ANTON 
BRUUN cruise 4B, sta. 211A, 16 Nov. 1963. SU 14666 (1, 51.6), 
Puri Beach, Orissa, 20 Mar. 1909. SU 14670 (2, 75.3-83.8), 
Ennur Fisheries Station, Madras, A. W. Herre, Jan. 1941. 
Additional material: SU 14667 (1), SU 14668 (6), SU 14669 
(3), USNM 218445 (5), USNM 218446 (1), USNM 218447 (2), 
CAS 33954 (43), FMNH 75583 (1), ZSI 3421/1 (2), ZSI 2347/ 
2 (2). 

Singapore: SU 30871 (1, 59.1), A. W. Herre, Mar. 1934. 
Additional material: SU 39470 (4). 

Indonesia: RMNH 5920 (3, 52.7-67.0, syntypes of M. oxy- 
cephalus), Sumatra and Amboina. Additional material: CAS 
36065 (3). 

Gulf of Thailand: CAS 13902 (3, 73.5-81.8), about 25 miles 
(40 km) offshore, 11°58’-12°05S’N, 99°56'30’—100°00'30"E, 18 
m, trawl, sandy bottom, 11-15 Mar. 1961. CAS 13903 (4, 64.7— 
72.2), 15-30 miles (24-48 km) offshore, 12°12'N, 100°14’— 
30’E, 26-27 m, trawl, sand and mud bottom, 23-25 July 1960. 
CAS 13906 (5, 55.5-79.4), 11°25'-45'30"N, 99°43’—53’E, 12-22 
m, trawl, mud and sand bottom, 6-11 May 1961. CAS 13907 
(4, 15.7 and 50.2-76.9), 13°21'45"N, 100°32'43”E, 14 m, trawl, 
mud bottom, 14 Dec. 1960. CAS 13910 (10, 43.9-73.9) and 
CAS 13908 (2, 67-72, cleared and stained), 11°40’-51'N, 
100°34'-39’E, 40-42 m, trawl, sand and shell bottom, 10-16 
Jan. 1961. Additional material: CAS 13913 (2), CAS 13914 (4), 
CAS 13916 (1), CAS 13948 (1), CAS 13949 (1), CAS 13950 (1), 
CAS 13951 (1), CAS 14490 (4), CAS 14495 (2), and CAS 14498 
(4, cleared and stained). 

Hong Kong: CAS 13911 (1, 48.4), 22°24'30"N, 113°53'30"E, 
15 m, trawl, mud bottom, 14 Aug. 1958. SU 61474 (1, 85.1), 
22°24'N, 114°25’E, 22 m, R. L. Bolin, 7-8 Jan. 1958. 

Mainland China: BMNH 1848.3.16.37 (1, 48.4, holotype of 
M. adamsii), China Seas, Belcher, no other data. SU 32941 
(1, 84.7), Tinghai, Chusan Is., Chekiang Province, A. W. 
Herre, May 1937. Additional material: SU 28177 (2). 

Taiwan: CAS 13952 (10), CAS 27740 (9), CAS 30002 (3), 
CAS 30310 (40), and SU 49444 (1). 

Japan: CAS 11243 (2, 76.2-87.0), Kobe, Settsu, D. S. Jor- 
dan and J. O. Snyder. CAS 11244 (3, 50.6-69.7), Onomichi, 
Bingo, D. S. Jordan and J. O. Snyder. SU 7370 (1, 86.3, ho- 
lotype of M. echigonius), Niigata, Echigo, D. S. Jordan and 
J. O. Snyder. USNM 55615 (1, 103, holotype of L. satsumae), 
Kagoshima, H. M. Smith, 16 June 1903. Additional material: 
SU 7173 (23), UMMZ 191787 (1), and UT 41826 (1), 51122 (1), 
51341 (1), 51408 (1), 51611 (1), 51612 (1), 51698(1), 51701 (1). 


462 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


DESCRIPTION (See also generic diagnosis; 
Figs. 2b, 3c; Tables 1— 2).—Dorsal fin normally 
with 9-11 (usually 10) spines and 10-12 (usually 
11) soft rays; total dorsal spines and rays 19-21 
(usually 21); dorsal spines sharp and strong; first 
dorsal spine equal to or longer than second, well 
separated from second. Anal fin with 2 spines 
and 7-10 (usually 9) soft rays; spines difficult to 
distinguish from soft rays; total anal fin elements 
9-12 (usually 11). Vertebrae 25 (1 specimen) or 
26 (14). Gill rakers total 11-16 on outside of first 
arch, 3-4 on upper arch, 8-12 on lower arch. 
Lateral line pores about 18-20. Gas bladder 
present. Pectoral fin reaching to middle of anal 
fin as maximum. Head spines well developed. 
Lachrymal bone with 2 spines over maxillary, 
first pointing down and forward, second down 
and back; second spine at least twice length of 
first and slightly bayonet shaped. 

General body shape and preserved coloration 
in Figure 3c, Color variable; dorsal portion of 
body with pale bars and stripes; ventral surface 
usually pale, without markings. Dorsal fin mem- 
branes dark distally. Anterior portion of soft 
dorsal fin with a large black area or spot, usually 
followed by oblique pale bars. Inside of pectoral 
fin and pectoral fin axil pale, without distinctive 
markings; outside of pectoral, pelvic, and anal 
fins blackish distally. Caudal peduncle crossed 
by a dark band, followed by 2 broad vertical 
bars across caudal fin. 

Measurements for 10 specimens (50.2- 
79.4 mm SL) in percent standard length: head 
36-44; orbit 10-12; snout 13-16; interorbital 
width 7-9; postorbital 15-19; pectoral 36-45; 
first dorsal spine 7-12; second dorsal spine 7- 
ll. 

DISTRIBUTION.—Minous monodactylus is a 
shallow-water species occurring from near shore 
to about 55 m. It is known from the western 
Indian Ocean and Red Sea to Indonesia and Ja- 
pan. It is the most widespread species of the 
genus. 


Minous pusillus Temminck and Schlegel 
(Figures 1, 4; Tables 1-2) 


Minous pusillus TEMMINCK AND SCHLEGEL, 1843:50 (original 
description; type locality Nagasaki, Japan). GUNTHER 
1860:149 (compiled). STEINDACHNER AND DODERLEIN 
1884:197 (description; Kagoshima Bay, Japan). MATSU- 
BARA 1943:409-412, fig. 140 (good description; compari- 
sons; variability; internal features; specimens from Japan). 
BOESEMAN 1947:60 (Von Siebold specimens as types of M. 
pusillus; lectotype designation). KAMOHARA 1952:68 (listed: 


Japan); 1958a:57 (Japan; compiled distribution); 1964:75 
(Japan; compiled distribution). 

Aploactis pusillus: BLEEKER 1859:6—7 (description; 2 speci- 
mens from Japan). 

Minaus pusillus: BLEEKER 1879c:12 (misspelled generic name; 
listed; Japan). 

Decterias pusillus: JORDAN AND STARKS 1904:154—-155 (as 
type of new genus Decterias: description; good figure; Jap- 
anese specimens). SMITH AND Pope 1906:483 (specimens 
from Susaki and Kagoshima, Japan). JoRDAN AND SEALE 
1906:38 (2 specimens from Panay, Philippines). ?FRANZ 
1910:74 (2 specimens; Japan; specimens with hydroids on 
body). JORDAN AND RICHARDSON 1910:52 (listed; Philip- 
pines). JORDAN, TANAKA, AND SNYDER 1913:247, fig. 182 
(listed; Japan; figure from Jordan and Starks 1904). 
SCHMIDT 1931:111 (3 specimens from Nagasaki, Japan). 
TANAKA 1931:36 (listed; Japan). RoxAS AND MARTIN 
1937:180 (listed; Panay, Philippines). HERRE 1951:473-474 
(synonymy; description; range; Philippine localities; com- 
pared with Japanese specimens); 1953:575 (listed; Philip- 
pines). 


MATERIAL EXAMINED.—Japan: RMNH 71]13a (1, 43.5 mm 
SL, lectotype of M. pusillus) and RMNH 713b-1 (11, 32-45, 
paralectotypes of M. pusillus), Von Siebold collection. SU 
7397 (11, 32.4-53.2), Wakanoura, Jordan and Snyder. 

East China Sea: CAS 27742 (47, 24-48), N of Taiwan, about 
26°-27°N, 121°-122°E, trawled, F. B. Steiner, 15 Oct. 1972. 

Taiwan Strait, collected by F. B. Steiner: CAS 13863 (12, 
33.8-51.1), 25°N, 120°E, in 90 m, Apr. 1971. CAS 13865 (1, 
47.8), 27°30'N, 121°30’E, in 80-100 m, 17 June 1971. CAS 
13866 (14, 35.6—53.8), 26°N, 121°E, in about 90 m, trawled, 16 
June 1971. CAS 13867 (6, 35.1-46.0), between Pescadores Is. 
and Taiwan, in 55-73 m, 7 Apr. 1971. CAS 14492 (9, 26.0- 
43.0) and CAS 14493 (36.5—48.2, cleared and stained), trawled 
in about 60 m, Feb. 1972. Additional material: CAS 15615 
(37), CAS 28128 (15), CAS 28181 (15), CAS 30311 (5), CAS 
30582 (1), CAS 34215 (1). 

Hong Kong: CAS 13868 (1, 50.7), off Lema I., 21°57’N, 
114°20’E, in 38-40 m, R. L. Bolin, 25 July 1958. SU 27995 (5, 
30.4-40.9), A. W. Herre, Oct. 1931. 

Philippines: CAS 29371 (1, 38), Mindanao, Agusan Proy- 
ince, Buena Vista, F. B. Steiner, 14 Apr. 1973. CAS 29407 
(1, 48.0), Mindanao, Agusan Province, Nasipit, F. B. Steiner, 
15 Apr. 1973. 


DESCRIPTION (See also generic diagnosis, Fig. 
4; Tables 1—2).—Dorsal fin with 9-11 (usually 10 
or 11) spines and 9-11 (usually 10) soft rays; 
total dorsal spines and rays 19-21 (usually 21 or 
20); dorsal spines weak and hairlike, difficult to 
distinguish from soft rays; first dorsal spine usu- 
ally less than % of second (0.5—0.75 length of 
second), and close together. Anal fin with 2 
spines and 8-9 (usually 8) soft rays; spines dif- 
ficult to distinguish from soft rays; total anal fin 
elements 10-11 (usually 10). Vertebrae 25 (16 
specimens). Gill rakers total 14-17 on first arch, 
3—4 on upper arch, 10-14 on lower arch. Lateral 
line pores 14-18. Gas bladder absent. Pectoral 
fin reaching to middle of anal fin as maximum. 


ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 463 


FIGURE 4. 
dusky pigment.) 


Head spines poorly developed; lachrymal bone 
with 2 short spines over maxillary, first pointing 
forward and down, second down and back; sec- 
ond spine slightly longer than first. 

General body shape and preserved coloration 
in Figure 4. Dorsal portion of body brownish or 
grayish, marbled slightly with paler areas. Ven- 
tral portion of body white, without markings. 
Dorsal fin membranes between anterior spines 
blackish distally. Outside of pectoral fin black; 
axil of pectoral fin grayish, fin becoming black- 
ish distally. Anal and pelvic fins dusky. Caudal 
fin rays with alternating dark and pale color, giv- 
ing effect of several narrow vertical dark bars. 

Measurements for 6 specimens (47.9-54.7 mm 
SL) in percent standard length: head 41-44; or- 
bit 12-14; snout 12-13; interorbital width 6-8; 
postorbital 16-18; pectoral fin 37-45; first dorsal 
spine 6-11; second dorsal spine 14-18. 

DISTRIBUTION.—Minous pusillus is a small 
species occurring from Japan south to Hong 
Kong and the Philippines. Depths of capture 
range from about 30 to 110 m. 


Minous inermis Alcock 
(Figures 1, 5; Tables 1-2) 


Minous inermis ALCOCK, 1889:299-300, fig. 4 on pl. XXII 
(original description; type locality Bay of Bengal, E of Sac- 
ramento Shoal on Godavari coast, in 70 fms [128 m]; 2 syn- 
types as ZSI 12444 and 12445); ALcock 1892:210-212 (sym- 
biosis between hydroid and M. inermis; additional 


Minous pusillus, about 41 mm SL, Japan. (From Matsubara, 1943, fig. 140a, but modified by addition of more 


specimens from India); ALCock 1894:116 (specimens from 
off Madras, Bay of Bengal, in 133 fms [243 m]; commensal 
hydroid on body). GOODE AND BEAN 1896:524 (Bay of Ben- 
gal, 70-150 fms [128-274m]; compiled). ALcock 1896:314 
(compiled); 1898:fig. 3 on pl. 18 (good illustration); 1899:30— 
31 (specimens from off Coromandel and Malabar coasts, 
India; depths, description). REGAN 1905b:329, 331 (listed; 
Gulf of Oman). NORMAN 1939:96 (listed specimens from 
Gulf of Aden, JOHN Murray sta. 37 and 194). SMITH 
1958:175, fig. m on pl. 8 (compiled). 

Minous longimanus REGAN, 1908:236—237, fig. 2 on pl. 28 
(original description; type locality Saya de Malha Bank, in 
47 fms [86 m]; holotype BMNH 1908.3.23.178). SMITH 
1958:175, fig. j on pl. 8 (compiled from Regan). 

Minous longipinnis LLoypD 1909:162, fig. 3 on pl. 47 (original 
description; type locality Gulf of Oman, 230 fms [420 m]; 
INVESTIGATOR Sta. 341; compared with M. inermis). SMITH 
1958:175, fig. i on pl. 8 (compiled from Lloyd; suggested 
may be synonym of M. longimanus). 

Paraminous inermis: FOWLER 1943:68 (placed in genus Par- 
aminous; compared with Paraminous quincarinatus). 


NOMENCLATURAL REMARKS.—Minous iner- 
mis as treated by Matsubara (1943:406—409, fig. 
139) is referable to M. quincarinatus Fowler, a 
species that was described in the same year as 
Matsubara’s treatment of the Japanese fauna. 
Other identifications of Japanese specimens 
(e.g., Franz, Kamohara) are referred question- 
ably to M. quincarinatus. Minous inermis A\l- 
cock was not treated by de Beaufort (in Weber 
and de Beaufort 1962) in his treatment of Indo- 
Australian fishes. Minous inermis of Herre (1951: 
475-476) is referred to M. trachycephalus 
(Bleeker). 


MATERIAL EXAMINED.—Burma: SU 14658 (1, 51.0), Mer- 
gui Archipelago, in 119 m, R.I.M.S. INVESTIGATOR sta. 535, 
17 Apr. 1913. 

Bay of Bengal: SU 67200 (1, 38.2), mouth of Hughli River, 
Lower Bengal, Sandheads, A. W. Herre, 6 Jan. 1928. ZSI 
12444 (1, 47.0, syntype of M. inermis) and ZSI 12445 (1, 48, 
syntype of M. inermis), Godarvari coast, Sacramento Shoal, 
in 128 m [both syntypes now in poor condition; no counts or 
measurements taken)). 

Arabian Sea area: ZSI 1158/1 (1, 72.6, syntype of M. lon- 
gipinnis), and ZSI 1159/1 (1, 73.9, syntype of M. longipinnis), 
Gulf of Oman, in 420 m, R.I.M.S. INVESTIGATOR sta. 341. 
BMNH 1908.3.23.178 (1, about 67, holotype of M. longiman- 
us), Saya de Malha Bank, 86 m, Gardiner Expedition. CAS 
13859 (18, 35.2-75.7), CAS 13860 (2, 65-79, cleared and 
stained), and ZSI F7402/2 (3, 42.0-67.0), western India, 
17°25'-21'N, 71°39'-41'E, 96-106 m, ANTON BRUUN cruise 
4B, sta. 202A, 13 Nov. 1963. CAS 13861 (1, 102), Somali 
coast, 11°04’N, 51°1S’E, 76-80 m, ANTON BRUUN cruise 9, 
sta. 451, 17 Dec. 1964. CAS 13862 (1, 61.0), 25°00’-24°59'N, 
63°30'-33'E, 35-37 m, ANTON BRUUN cruise 4B, sta. 242A, 
27 Nov. 1963. CAS 14496 (3, 42.8-61.5), western India, 
21°11'-08'’N, 69°16’— 13’E, 70-72 m, ANTON BRUUN cruise 
4B, sta. 213A, 17 Nov. 1963. USNM 218449 (8, 35.0-65.8), 
western India, 17°41-45'N,71°33'—32’E, 90 m, ANTON BRUUN 
cruise 4B, sta. 202B, 14 Nov. 1963. USNM 218450 (2, 65.7- 
66.4), western India, 21°52’-55'N, 68°06’E, 115-117 m, An- 
TON BRUUN cruise 4B, sta. 219A, 18 Nov. 1963. 


DESCRIPTION (See also generic diagnosis; Fig. 
5; Tables 1-2).—Dorsal fin with 9-10 spines and 
12-14 (usually 13-14) soft rays; total dorsal fin 
spines and rays 22-24 (usually 23); dorsal spines 
thin and flexible; first dorsal spine short, usually 
less than % (0.2—0.4) of second and close to base 
of second. Anal fin with 2 spines and 8-11 (usu- 
ally 10) soft rays; spines difficult to distinguish 
from soft rays; total anal fin elements 10-13 
(usually 11-12). Vertebrae 26 (3 specimens) or 
27 (16). Gill rakers total 14-18 on outside of first 
arch, 4 on upper arch, 10-14 on lower arch. Lat- 
eral line pores 16-18. Gas bladder present. Pec- 
toral fin very long, reaching to over end of anal 
fin. Head spines moderately developed. Lach- 
rymal bone with two sharp spines over maxil- 
lary; first points down and forward, second 
down and slightly back; spines about equal in 
length. 

General body shape and preserved coloration 
as in Figure 5. First 5—6 dorsal fin spines with 
dark skin flap at end. Body coloration variable. 
Smaller specimens with pale spots on a light 
background on the sides of body; spots not pres- 
ent on one larger specimen. Ventral portion of 
body pale, lacking markings. Anal fin black dis- 
tally. Pectoral fin base light brown, the fin in- 
creasing in darkness distally; inside of pectoral 
fin light gray; axil with white spots on a gray 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


background, uniform gray distally. Pelvic fins 
with pale spots on a gray background. Caudal 
fin mostly pale, dark distally. 

Measurements for 16 specimens (52.9-102 
mm SL) in percent of standard length: head 41- 
48; orbit 10-13; snout 14-15; interorbital width 
6-7; postorbital 19-23; pectoral fin 50-60; first 
dorsal spine 4—9; second dorsal spine 11-18. 

DISTRIBUTION.—Minous inermis is known 
from the northern Indian Ocean in coastal areas 
from the Bay of Bengal west to the Gulf of 
Oman, Somalia, and from the Saya de Malha 
Bank. Depths of capture range from 35 to 420 
m. 


Minous trachycephalus (Bleeker) 
(Figures 1, 6; Tables 1-2) 


Aploactis trachycephalus BLEEKER 1854b:449, 451-452 (orig- 
inal description; type locality Celebes, Dutch East Indies). 

Corythobatus trachycephalus: BLEEKER 1865: 282 [286] (list- 
ed). 

Minous inermis (not of Alcock): HERRE 1951:475—476 (10 
specimens from Manila Bay, Luzon, 17-25 fms [31-46 m]; 
description; provisional identification). 

Minous trachycephalus: GUNTHER 1860:149 (compiled). 
BLEEKER 1876b:9-13, 61-63, fig. 3 on pl. 2 (description; 
good figure; specimens from Nias, Celebes, and Amboina); 
1879a:fig. 4 on pl. 415 (figure from Bleeker 1876b). FOWLER 
AND BEAN 1922:63 (Takao, Formosa; brief description). 
HERRE 1936:367 (1 from Subic Bay, Philippines). Roxas 
AND MARTIN 1937:181 (compiled). HERRE 1951:477-478 
(compiled). DE BEAUFORT in WEBER AND DE BEAUFORT 
1962:108-109 (wrongly included M. superciliosus in syn- 
onymy; description). 


NOMENCLATURAL REMARKS.—Bleeker 
(1854b) mentions one specimen of 75 mm in 
length from the type locality Celebes. This spec- 
imen is apparently now mixed in RMNH 5901 
containing five specimens. Two of these speci- 
mens are of about 75 mm in total length, and 
presumably one is the type. 

The single specimen from Japan identified as 
this species by Matsubara (1943) seems referable 
to M. pictus as do some subsequent listings of 
trachycephalus from Japan (e.g., Kamohara 
1958a, 1960, 1964). The specimen from the Mal- 
abar Coast of India that Alcock (1896) identified 
as trachycephalus is M. coccineus. 


MATERIAL EXAMINED.—Ceylon: USNM 218448 (1, 35.0), 
Gulf of Mannar, 8°39’—53’'N, 79°37'-46'E, 11-18 m, T. R. Rob- 
erts, 3-7 Apr. 1970. 

Gulf of Thailand: CAS 13871 (1, 48.9), 12°19'15’N, 
100°43'40"E, 33m, 13 Dec. 1960. CAS 13872 (1, 53.4), 12°12'N, 
100°17'E, about 28 m, 8-10 Sep. 1960. CAS 13873 (1, 42.1), 
12°19’15"N, 100°43’40’E, in 33 m, 18 Dec. 1960. CAS 13874 
(2, 46.9-56.3), about 12°23’N, 100°33’E, about 36 m, 17-21 


ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 465 


FIGURE 5. 


Dec. 1960. CAS 13875 (2, 56.9-57.0), and CAS 14494 (1, 60, 
cleared and stained), 11°40’—S1’N, 100°34’—39’E, about 40-42 
m, 10-16 Jan. 1961. CAS 13876 (1, 64.8), 12°08’-20'N, about 
100°14'—22'E, about 26-28 m, 30 July—1 Aug. 1960. CAS 13877 
(1, 65.7) and CAS 13878 (1, 57.9, cleared and stained), 12°05’— 
19’N, 100°21'— 41’E, 27-35 m, 2-6 Apr. 1961. 

Viet Nam: CAS 13870 (1, 35.3), 12°09'30"N, 109°14’'15"E, 
15 m, R. L. Bolin, 23 Feb. 1960. 

Philippines: SU 9664 (2, 30.0-36.1), Iloilo I., Panay, G. A. 
Lung. SU 29784 (1, 54.6), Subic Bay, Zambales Prov., J. C. 
Thompson, 1906. UWCF 7244 (9, 31.5—39.0), Manila Bay, 
Luzon, A. W. Herre, 25 May 1949. Additional material: 
USNM 168176 (3). 

Taiwan: USNM 76650 (1). 

Celebes: RMNH 5901 (5), see Nomenclatural Remarks. 


DESCRIPTION (See also generic diagnosis; Fig. 
6; Tables 1—2).—Dorsal fin with 10-11 (usually 
11) spines and 8—10 (usually 9-10) soft rays; total 
dorsal spines and rays 19-21 (usually 20-21); 
dorsal spines moderately strong, thin, and flex- 
ible in specimens under about 40 mm SL; first 
dorsal spine short, usually less than half (0.3- 
0.5) length of second and close at base to sec- 
ond. Anal fin with 2 spines and 7-9 (usually 8) 
soft rays; spines difficult to distinguish from soft 
rays; total anal fin elements 9-11 (usually 10). 
Vertebrae 24 (1 specimen) or 25 (6). Gill rakers 
total 10-13 on outside of first arch, 24 on upper 
arch, 7-10 on lower arch. Lateral line pores 15- 
16. Gas bladder absent. Pectoral fin reaching to 
middle of anal fin as a maximum. Head spines 
poorly developed. Lachrymal bone with 2 short 
spines over maxillary, first points down and for- 


Minous inermis, ZSI 1158—9/1, about 73 mm SL, Gulf of Oman. (From Lloyd 1909; fig. 3 on pl. 47.) 


ward, second down and back; second spine 
longer than first. 

General body shape and preserved coloration 
as in Figure 6. Dorsal half of body grayish 
brown. Underside white, without markings. 
Body and fins with scattered dark spots. Soft 
dorsal fin pale anteriorly, becoming darker dis- 
tally and posteriorly. Pectoral fin base pale; pec- 
toral fin mostly black. Inside of pectoral fin 
mostly gray; axil pale, with many large white 
spots forming a hexagonal pattern; fin gray dis- 
tally, the rays occasionally with minute dark 
black spots. Pelvic fins gray, sometimes with 
black spots on rays. Caudal fin rays with alter- 
nating gray and pale color; fin membranes most- 
ly clear, with many minute dark flecks. 

Measurements for 14 specimens (35.0-65.7 
mm SL) in percent standard length: head 43-48; 
orbit 12-16; snout 14-18; interorbital width 7-9; 
postorbital 14-18; pectoral fin 40-46; first dorsal 
spine 3—5; second dorsal spine 11-14. 

DISTRIBUTION.—Minous trachycephalus is 
known from areas bordering the South China 
Sea and from northern Indonesia and Ceylon. A 
wider distribution is expected. This species ap- 
pears to be one of the more shallower-living spe- 
cies. Few depths of capture are available, but 
are between 11-46 m. 


Minous coccineus Alcock 
(Figures 1, 2d, 7; Tables 1-2) 


Minous coccineus ALCOCK 1890:428—429 (original description; 


466 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


FIGURE 6. 


type locality India, Ganjam Coast, 28-30 fms [51-54 ml], 
sand and shell bottom). 

Minous superciliosus GILCHRIST AND THOMPSON 1908:177- 
178 (original description; type locality NW 7.5 miles [12 km] 
from Amatikulu Conical Hill, PrleTER FAURE, shrimp trawl, 
26 fms [47 m]); 1917:410 (compiled). BARNARD 1927:921 
(description; Zululand coast, 26 fms [47 m]; close to M. 
trachycephalus); 1947:199, fig. 1 on pl. 24 (brief description; 
figure poor). FOWLER 1934:486 (description; distinct from 
M. monodactylus and M. trachycephalus). VON BONDE 
1924:3, 29-30 (specimens from PICKLE stations 169, 171, 
179, and 193, in 27-29 fms [49-53 m]). SMITH 1958:175—176, 
fig. H on pl. 8 (description; synonymy; specimen from off 
Durban in 40 fms [73 m]). 

Minous trachycephalus: ALCOCK 1896:313 (misidentified; list- 
ed from Malabar Coast, India; R.I.M.S.INVESTIGATOR col- 
lection). SMITH 1949:375, fig. 1054 (misidentified; brief de- 
scription; thought M. superciliosus was a synonym of 
trachycephalus). DE BEAUFORT in WEBER AND DE BEAU- 
FORT 1962:109—-110 (in part; wrongly included references to 
M. superciliosus in synonymy of M. trachycephalus). 


MATERIAL EXAMINED.—South Africa: ANSP 77866 (1, 
76.0), Natal, PickLeE Survey, H. W. Bell Marley, 17 Aug. 
1930. 

Red Sea: CAS 14489 (31, 53.0-80.0) and USNM 218451 (1, 
63.0), Bay of Massawa, about 15°40'N, 39°36’E, 79 m, L. 
Knapp, 20 Sep. 1971. USNM 218452 (1, 62.0), off Ajuz, about 
16 miles (26 km) S of Massawa, 15°18’N, 40°17’E, 29-31 m, 
mud bottom, L. Knapp, 19 Sep. 1971. USNM 218453 (1, 64), 
Massawa Bay, 15°40'N, 39°36’E, 79 m, L. Knapp, 20 Sep. 
1971. USNM 218454 (3, 66-69), Massawa Bay, 15°40’N, 
40°23’E, 79 m, L. Knapp, 21 Sep. 1971. 

Arabian Sea: ZSI 13760 (1, 85.0), Malabar Coast, 81 m, 
western India, R.I.M.S. INVESTIGATOR collection. 


Minous trachycephalus, CAS 13876, 64.8 mm SL, Gulf of Thailand. 


Bay of Bengal: ZSI 12924 (1, 77.0, syntype of M. cocci- 
neus), off Ganjam Coast, 51 m, R.I.M.S. INVESTIGATOR, 17— 
21 Feb. 1890. BMNH 1978.4.19.3 (1, 48.0) and ZSI F7403/2 
(1, 41.2), Bangladesh, 21°00’N, 91°59’'E, 23-25 m, ANTON 
BRUUN cruise 1, sta. 46, 5 Apr. 1963. SU 67169 (3, 47.4-86.3) 
and SU 67170 (2, 60.3-74.9, cleared and stained), India, mouth 
of Hughli River, Sandheads, A. W. Herre, 6 Jan. 1928. ZSI 
2485/1 (2, 74-76), Ganjam Coast. Additional material: ZSI 
Madras collections from Kakinda and Madras, India. 

Burma: USNM 218455 (2, 58.6—75.3), Mergui Archipelago, 
9°54'N, 97°42'E, 70 m, ANTON BRUUN cruise 1, hydro sta. 21, 
24 Mar. 1963. 

Java: SU 49320 (1, 69.8), no other data. 

Thailand: CAS 13893 (1, 78.9), 12°26’-27'30’N, 
26'E, 26 Feb.—2 Mar. 1961. 

Taiwan: SU 49442 (1, 80), no other data. 


101°20'- 


DESCRIPTION (See also generic diagnosis; 
Figs. 2d, 7; Tables 1—2).—Dorsal fin with 10-11 
(usually 11) spines and 11-12 (usually 12) soft 
rays; total dorsal spines plus rays 21—23 (usually 
23); dorsal spines sharp and moderately strong; 
first dorsal spine short, sometimes hidden under 
skin, less than half (0.2—-0.4) length of second 
spine and close to second spine at base. Anal fin 
with 2 spines and 9-10 (usually 10) soft rays; 
spines difficult to distinguish from soft rays; to- 
tal anal fin elements 11—12 (usually 12). Verte- 
brae 26 (7 specimens). Gill rakers total 11-14 on 
outside of first arch, 3-4 on upper arch, 7—10 on 
lower arch. Lateral line pores 18-19. Gas blad- 
der present. Pectoral fin reaching to middle of 


ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 467 


ge ere = oA 
aes ater 
AN 
aah 


FiGurE 7. Minous coccineus, about 63 mm SL 


anal fin as a maximum. Head spines well devel- 
oped. Lachrymal bone with 2 sharp spines over 
maxillary, first points down and forward, second 
down and to rear; first spine about half length 
of second. 

General body shape and preserved coloration 
as in Figure 7. Body coloration variable. Pos- 
terior part of spinous dorsal fin and soft dorsal 
fin usually with alternating white and brown 
bands extending down and back, often onto the 
upper surface of the body. Lower portion of 
body pale, without markings. Pectoral fin base 
light brown, with color from inside fin showing 
through; axil pale; inside of pectoral fin with dis- 
tinct black spots on a pale background (Fig. 2d). 
Pelvic and anal fins dark. Caudal fin pale, with- 
out markings. 

Measurements for 9 specimens (48.2—86.3 mm 
SL) in percent of standard length: head 39-43; 
orbit 12-14, snout 13-14; interorbital width 9- 
10; postorbital 16-19; pectoral fin 33-41; first 
dorsal spine 4—5; second dorsal spine 14-17. 

DISTRIBUTION.—Minous coccineus is known 
from coastal waters of South Africa, the Red 
Sea, Arabian Sea, Bay of Bengal, Burma, Gulf 
of Thailand, Java, and Taiwan in depths ranging 
from 23 to 81 m. 


Minous pictus Gunther 
(Figures 1, 8a; Tables 1-2) 


Minous pictus GUNTHER 1880:41, pl. 18, fig. D (original de- 
scription; type locality Arafura Sea, S of New Guinea, 


, South Africa. (From Smith 1949; fig. 1054, p. 375.) 


9°59'S, 139°42'E, CHALLENGER sta. 188). JORDAN AND 
SEALE 1906:378 (listed; New Guinea). FOWLER 1928:299 
(compiled). DE BEAUFORT in WEBER AND DE BEAUFORT 
1962:111 (redescribed type). 

Minous trachycephalus: MATSUBARA 1943:405—406 (misiden- 
tified; 1 specimen from Japan; description). KAMOHARA 
1958a:76 (listed; Japan); 1960:27 (compiled); 1964:75 (com- 
piled). 


MATERIAL EXAMINED.—New Guinea: BMNH 1879.5.14.371 
(1, 46.1, holotype of M. pictus), Arafura Sea, S of New 
Guinea, 9°59'S, 139° 42'E, CHALLENGER Sta. 188, 56 m. 

Philippines: SU 39136 (1, 41.1), Manila Bay, Luzon, A. W. 
Herre, 7 June 1940. USNM 218444 (2, 66.0-76.3), Manila Bay, 
purchased at fish market, F. Schwartz, 13 May 1939. Addi- 
tional material: CAS 32650 (2), 32770 (1), 32943 (1), 33090 (1), 
and USNM 99787 (6). 

Taiwan: CAS 13864 (1, 45.5, cleared and stained) and CAS 
13901 (2, 72.8-79.7), 25°N, 120°E, 90 m, trawl, muddy bot- 
tom, F. B. Steiner, Apr. 1971. Additional material: CAS 14491 
(6), CAS 14497 (6), CAS 28180 (24), CAS 29975 (5), CAS 30309 
(1), CAS 34214 (2), SU 49443 (1), USNM 200233 (2). 

Hong Kong area: CAS 13869 (1, 78.1), off Lema Is., 
21°57'N, 114° 20’E, 38-40 m, shrimp trawl, ALISTER HARDY, 
R. L. Bolin, 25 July 1958. CAS 13894 (2, 73.2-85.1) and CAS 
13895 (1, 76, cleared and stained), 21°35'N, 114°00’E, 42 m, 
mud bottom, R. L. Bolin, 24 July 1958. CAS 13896 (2, 65.5— 
78.5), 22°06'30"N, 114°16'30"E, 33 m, sticky gray mud, ALIs- 
TER Harpy, R. L. Bolin, 29 Dec. 1957. CAS 13900 (1, 120), 
about 150 miles (ca. 240 km) west of Hong Kong, F. B. Stei- 
ner, 14 June 1971. CAS 14487 (1, 63.5), South China Sea, 
20°32'N, 112°51’E, R. L. Bolin, 23 July 1958. SU 60895 (3, 
58.4-68.8), 21°56’N, 114°23’E, R. L. Bolin, 28 Dec. 1957. SU 
61010 (4, 67.7-68.2), Mira Bay, 22°28’N, 114°23’E, R. L. Bolin 
and party, 7 Jan. 1958. Additional material: CAS 14488 (2), 
CAS 27741 (10). 

Viet Nam: CAS 13897 (1, 54.1), 15°41'30’N, 108°42’E, fine 
green mud, M/V STRANGER, R. L. Bolin, 27 Feb. 1960. Ad- 
ditional material: CAS 13898 (2). 


468 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


FIGURE 8. 
off western India. 


Indonesia: CAS 41051 (1, 32.0), Java Sea, 5°58’S, 106°48’E, 
24 m, muddy, F. B. Steiner, 5 Dec. 1975. 

DESCRIPTION (See also generic diagnosis; Fig. 
8a; Tables 1-2).—Dorsal fin with 10-12 (usually 
11) spines and 11-13 (usually 12) soft rays; total 
dorsal spines plus soft rays 22-24 (usually 23); 
dorsal spines sharp and moderately strong; first 
dorsal spine short, usually less than one fourth 
(0.2-0.4) of second and close to base of second. 
Anal fin with 2 spines and 9-11 (usually 10) soft 
rays; spines difficult to distinguish from soft 
rays; total anal fin elements 11—13 (usually 12). 
Vertebrae 24 (1 specimen), 25 (2), 26 (8), 27 (1). 
Gill rakers total 13-17 on first gill arch, 3-5 on 
upper arch, 9-12 on lower arch. Lateral line 
pores 18-20. Gas bladder present. Pectoral fin 
reaching to middle of anal fin as a maximum. 


(a) Minous pictus, CAS 28180, 125 mm SL, Taiwan. (b) M. dempsterae, holotype, USNM 218417, 84 mm SL, 


Head spines well developed. Lachrymal bone 
with two sharp spines over maxillary, first points 
down and forward, second down and to rear; 
second spine about twice length of first. 
General body shape and preserved coloration 
as in Figure 8a. Body coloration variable; alter- 
nating pale and dark oblique bands present to 
some degree in all specimens, these bands ex- 
tend obliquely back from under the posterior 
portion of the spinous dorsal fin and the soft 
dorsal fin. Undersurface of body pale, without 
markings. Anal fin pale, sometimes with outer 
margin tinged with black. Pectoral fin base light 
brown or pale; pectoral fin blackish. Inside of 
pectoral fin variable, axil light brown or pale, 
with gray stripes or rows of spots radiating out- 
ward along pectoral rays (similar to Fig. 2c). 


ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 


Pelvic fins brown, darker distally. Caudal fin 
pale, with a dark edge. 

Measurements for 16 specimens (46.1—85.1 n 
mm SL) in percent standard length: head 40-46; S 
orbit 12-16; snout 13-16; interorbital width 9- 
11; postorbital 16-19; pectoral fin 38-45; first 
dorsal spine 3-4; second dorsal spine 10-14. 

DISTRIBUTION.—Minous pictus is known 
from the Arafura Sea, Philippines, Taiwan, 
Hong Kong, and Viet Nam, in depths from 27- 
90 m. 


14508 


79.9 


CAS 
13891 
XI,12 


71.8 
XI,12 


Minous dempsterae Eschmeyer, Hallacher, and 
Rama-Rao, new species 
(Figures 1, 2e, 8b, 9; Tables 1-3) 


No literature applies to this species. 


CAS 
13887 
XI,11 


MATERIAL EXAMINED.—AIl type-specimens were collected 
by the International Indian Ocean Expedition, ANTON BRUUN 
cruise 4B, Nov. 1963. 

Holotype: USNM 218417 (84.0 mm SL), off western India, 
21°11'-08’N, 69°16’—13’E, 70-72 m, sta. 213A. 

Paratypes: Western India: BMNH 1978.4.19:1-2 (2, 66.6— 
83.1), CAS 13891 (3, 71.6-79.9), CAS 13892 (2, 74-77, cleared 
and stained), RUSI 953 (2, 62.8-73.2), USNM 218418 (2, 72.6— 
81.0), and ZSI F7401/2 (2, 69.4-70.0), all from 17°41’-45'N, 
71°33'-32'E, 90 m, sta. 202B. CAS 13884 (3, 75.0-88.6), 
21°52'—55’'N, 68°06’E, 115-117 m, brown mud, sta. 219A. CAS 
13887 (3, 71.8-83.2), 17°25’-21'N, 71°39’-41’E, 96-106 m, 
greenish sand and mud, sta. 202A. CAS 14508 (1, 75.1), 
20°49'—52’N, 69°41’-39’E, 60-62 m, sta. 209A. CAS 14509 (5, 
72.4-81.8) and USNM 218419 (4, 70.0-76.5), taken with the 
holotype. USNM 218420 (1, 59.6), 17°54'-57'N, 72°27'-23’E, 
46-55 m, sta. 201A. ANSP 136720 (2, 62.0-67.7) and USNM 
218421 (2, 69.7-70.1), 22°32’-31'N, 68°07’-05’E, 57 m, sta. 
221A. Pakistan: CAS 13883 (3, 60.1-84.9), 23°45'—43'N, 
67°23'—26'E, 23-24 m, greenish-brown clay bottom, sta. 228A. 
Gulf of Oman: CAS 13886 (1, 56.6), 26°10’-13'N, 57°02'E, 64— 
55 m, green mud, sta. 256A. 

Non-type material: CAS 29591 (38 specimens), India, Bom- 
bay, 5-10 m, trawl mud-sand bottom, F. B. Steiner, 4-10 Nov. 
1974. 


80.4 
XI,12 


CAS 

13884 
79.9 
x2 


88.6 
XI1,12 


84.0 
XI,12 


DESCRIPTION (See also generic diagnosis; 
Figs. 2e, 8b, 9; Tables 1—3).—Dorsal fin with 10- 
11 (usually 11) spines and 11-13 (usually 12) soft 
rays; total dorsal spines and rays 22—24 (usually 
23); dorsal spines sharp and moderately strong; 
first dorsal spine less than half (0.2—0.4) of sec- 
ond; close to base of second. Anal fin with 2 
spines and 9-11 (usually 9-10) soft rays; anal 
spines difficult to distinguish from soft rays; to- 
tal anal fin elements 11-13 (usually 11-12). Ver- 
tebrae 25 (1 specimen), 26 (18) and 27 (1). Gill 
rakers total 11-15 on outside of first arch, 3-5 
on upper arch, 8-10 on lower arch. Lateral line 
pores 18-20. Gas bladder present. Pectoral fin 
reaching to middle of anal fin as a maximum. 


TABLE 3. COUNTS AND MEASUREMENTS FOR SOME TYPE-SPECIMENS OF Minous dempsterae (measurements in mm, percent standard length in parentheses). 
Holotype 
USNM 


Standard length 
Dorsal rays 
Anal rays 


11,10 


ae il 


11,10 


il sp 


11,10 


il ap 


11,9 


Wl ae 1 


II,9 
Wil sel 


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8) 


II 
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Gill rakers 


ot 


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469 


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3.6(05) 
5.3(07) 


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11.3(15) 


EN SN ER, SS ES SS 
SC AS GN be Se 8 22 
SS Sa oS SS SS SS 
SOn= OOH At = 
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loa) _-— foal _ 


—- rT rTro~wWT~TrorereTwrrvrerae 


SES BONG EON See SCAG 
own tet Omran A + 
teem roa) _ 
en 
Yas Sows |e & 
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o 5 V9 
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moma Ee Re Se 
_— — 
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<2) 5S) 2S) fe) fe} fel el oS 
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470 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


FIGURE 9. 


Head spines well developed (Fig. 9). Lachrymal 
bone with 2 sharp spines over maxillary, first 
points down and forward, second points out and 
to rear; first spine about half length of second. 

General body shape and preserved coloration 
as in Figure 8b. Anterior spinous dorsal fin 
membranes blackish distally. Posterior spinous 
dorsal fin membranes and soft dorsal fin often 
with pale bands extending across fins and onto 
body; pale bands sometimes bordered with a 
thin dark line. Ventral part of body pale, without 
markings. Pectoral fin base pale, fin darker. In- 
side of pectoral fin as in Figure 2e, axil pale, 
inner surface with small irregular pale spots on 
a dark background. Pelvic and anal fins dusky. 
Caudal fins pale. 

Measurements for 10 specimens (71.8—-88.6 
mm SL) in percent standard length: head 43-47; 
orbit 12-13; snout 14-16; interorbital width 9- 
11; postorbital 18-22; pectoral fin 38—46; first 
dorsal spine 3—5; second dorsal spine 11-14. 

DISTRIBUTION.—Minous dempsterae is known 
from off western India, Pakistan, and the Gulf 
of Oman, in depths from 23 to 117 m, and from 
off Bombay, India, in 5-10 m. 

NAME.—The species is named for Lillian J. 
Dempster, a friend and colleague, in recognition 
for her assistance in the preparation of this and 
other papers on scorpionfish. 


Diagram of head spines in Minous dempsterae, based on specimens in CAS 14509. 


LITERATURE CITED 


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. 1890. Natural history notes from H.M. Indian Marine 

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1892. A case of commensalism between a gymno- 

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214, 1 fig. 

. 1894. Natural history notes from H.M. Indian Marine 

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Bay of Bengal and from the Laccadive Sea. J. Asiat. Soc. 

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. 1896. Natural history notes from H.M. Indian Marine 

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supplementary list of the marine fishes of India, with de- 

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Bengal 65 (Pt. 2, no. 3):301—338. 

. 1898. Illustrations of the zoology of the Royal Indian 

Marine Surveying Steamer “‘Investigator’’ .... Fishes- 

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. 1899. A descriptive catalogue of the Indian deep-sea 
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BARNARD, H. 1927. A monograph of the marine fishes of 
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18-37. 


ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 471 


. 1947. A pictorial guide to South African fishes, ma- 
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BLEEKER, P. 1849. Bijdrage tot de kennis der Scleroparei van 
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1851. Over eenige nieuwe soorten van Scleroparei 

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1:17-27. 

. 1852a. Bijdrage tot de kennis der ichthyologische fau- 

na van de Moluksche eilanden. Visschen van Amboina en 

Ceram. Natuurk. Tijdschr. Ned.-Indié 3:229-309. 

1852b. Nieuwe bijdrage tot de kennis der ichthyolo- 

gische fauna van het eiland Banka. Natuurk. Tijdschr. 

Ned.-Indié 3:715—738. 

. 1854a. Overzicht der ichthyologische fauna van Su- 

matra, met beschrijving van eenige nieuwe soorten. Na- 

tuurk. Tijdschr. Ned.-Indié 7:49-108. 

1854b. Zesde bijdrage tot de kennis der ichthyolo- 

gische fauna van Celebes. Natuurk. Tijdschr. Ned.-Indié 

7:449-452. 

1859. Vijfde bijdrage tot de kennis der ichthyolo- 

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$:1-12. 

. 1861. Mededeeling omtrent vischsoorten nieuw voor 

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Afd. Natuurk. 12(1):28-63. 

. 1865. Enumeration des espéces de poissons actuelle- 

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. 1874. Poissons de Madagascar. E. J. Brill, Leiden, 

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1879a. Atlas ichthyologique des Indes Orientales 

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CANTOR, T. 1849. Catalogue of Malayan fishes. J. Asiat. Soc. 
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, AND E. STECHOW. 1908. Symbiose zwischen einem 
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GILcHRIsT, J. D. F., AND W. W. THOMPSON. 1908. Descrip- 
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, AND . 1917. A catalogue of the sea fishes re- 
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Goope, G. B., AND T. H. BEAN. 1896. Oceanic ichthyology 
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GUNTHER, A. 1860. Catalogue of the acanthopterygian fishes 
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. 1880. Report on the shore fishes procured during the 
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Herre, A. W. C. T. 1932. Fishes from Kwangtung Province 
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. 1936. Notes on fishes in the Zoological Museum of 

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. 1951. A review of the scorpaenoid fishes of the Phil- 

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. 1953. Check-list of Philippine fishes. U.S. Fish Wildl. 

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, AND G. S. Myers. 1937. A contribution to the ich- 
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Husner, J. 1816. Verzeichniss bekannter Schmettlinge, 
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472 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 20 


JoRDAN, D. S., AND C. L. Huss. 1925. Record of fishes 
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, AND R. E. RICHARDSON. 1910. Check-list of the spe- 

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, AND A. SEALE. 1906. The fishes of Samoa... . Bull. 

Bur. Fish. 1905, 25:173-455, 111 text figs., pls. 38-53. 

, AND E. C. Starks. 1904. A review of the scorpenoid 

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, S. TANAKA, AND J. N. SNYDER. 1913. Catalogue of 

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KAMOHARA, T. 1930. Fishes collected in the vicinity of Kas- 
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1952. Revised description of the offshore bottom- 

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1958a. A catalogue of fishes of Kochi Prefecture 

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Kaup, J. J. 1858. Einiges uber die Acanthopterygiens a joue 
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MACHAN, B. 1930. Fische aus Padang. Ann. Naturh. Mus. 
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McCuttocn, A. R. 1915. Report on some fishes obtained by 
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ESCHMEYER, HALLACHER, & RAMA-RAO: SCORPIONFISH GENUS MINOUS 473 


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C253% 
NH 
PROCEEDINGS 
OF THE 
CALIFORNIA ACADEMY OF SCIENCES 
FOURTH SERIES 
Vol. XLI, No. 21, pp. 475-500; 10 figs., 1 table. January 24, 1979 


FISHES OF THE SCORPIONFISH SUBFAMILY CHORIDACTYLINAE 
FROM THE WESTERN PACIFIC AND THE INDIAN OCEAN 


By 
William N. Eschmeyer 
California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 


Kaza V. Rama-Rao 


Marine Biological Station, Zoological Survey of India 
100 Santhome High Road, Madras 600 028, India 


and 
Leon E. Hallacher’ 
California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 


ABSTRACT: The Indo-West Pacific scorpionfish subfamily Choridactylinae Kaup contains two genera and 
10 species. The subfamily synonyms include Pelorinae Gill and Inimicinae Gill, the latter being the subfamily 
name applied to this group until now. Two species, with three free lower pectoral rays, are placed in the 
genus Choridactylus Richardson, which includes the nominal genera Chorismodactylus Gunther and Chori- 
dactylodes Gilchrist. Eight species, with two free lower pectoral rays, are included in /nimicus Jordan and 
Starks; Pelor Cuvier, Simopias Gill, and Chorismopelor Chevey are synonyms of Inimicus. Three species 
groups are recognized in Inimicus: (1) I. filamentosus (Cuvier); (2) I. caledonicus (Sauvage), I. cuvieri 
(Gray), I. didactylus (Pallas), and J. sinensis (Valenciennes); (3) J. brachyrhynchus (Bleeker), J. japonicus 
(Cuvier), and J. joubini (Chevey). Nine other nominal species are recognized as synonyms, including /. 
bifilis Fowler and I. cirrhosus McKay. Salient features used in the descriptions of species and separation of 
species groups include fin-ray counts, elevation of the orbits, head shape, height of interspinous membranes, 
and color pattern on the inside of the pectoral fin. Descriptions and figures are provided for all species. 

The subfamily is distributed throughout the Indian Ocean and the western Pacific to Japan, the major 
islands of the Indo-Australian Archipelago to tropical Australia, and east to Melanesia. The species are 
bottom-dwelling fishes of silty and sandy substrates. They are nearshore or slightly offshore forms, most 
occurring shallower than 40 m. 

Certain morphological and behavioral features allow burying in the substrate, and these features are 
discussed and illustrated based on aquarium observations. 


INTRODUCTION and are characterized by having the lower two 


Scorpionfishes of the subfamily Choridactyli- oF three pectoral fin rays free. These rays are 


nae occur in the Indo-West Pacific faunal region used in certain forms of locomotion, and their 
morphology has been discussed by Samuel 


' Present address: 198 Park Avenue, Monterey, California (1961). Matsubara (1943) defined the subfamily 
93940. (as Pelorinae) on an osteological basis, including 
[475] 


476 


in the subfamily only the genus Inimicus. We 
refer to this subfamily the genera Choridactylus, 
Chorismopelor, and Simopias, the latter two 
being synonymous with Jnimicus. These genera 
were often placed in the family Synanceiidae by 
previous workers (Bleeker 1874, ‘‘Synanceioi- 
dei’; Herre 1951, ‘‘Synancetidae’’; Smith 1958, 
“*Synanciidae’’; de Beaufort 1962, ‘‘Synancei- 
dae’’). Reasons for recognizing the Choridac- 
tylinae [formerly Inimicinae], Minoinae, and 
Synanceinae as subfamilies of the Scorpaenidae 
are provided by Eschmeyer and Rama-Rao 
(1973). 

This paper is one of a continuing series de- 
voted to defining the species of the family Scor- 
paenidae and their allies. The species of the 
subfamily Choridactylinae are poorly known 
and no revision of the group is available. Earlier 
studies (e.g., Jordan and Starks 1904; Matsubara 
1943; Herre 1951; Smith 1958; de Beaufort in 
Weber and de Beaufort 1962) were of a regional 
nature, although McKay (1964), in describing a 
new species of the genus /nimicus, provides a 
brief discussion of most of the nominal species 
of the genus, as does Condé (1977). Condé also 
includes excellent color photographs of J. fila- 
mentosus and behavioral observations. 

The limited information available on the ecol- 
ogy of these fishes seems to indicate that they 
are bottom-dwelling, marine forms which tend 
to inhabit open sandy or silty substrates (McKay 
1964). The group is largely a coastal one, with 
a shallow bathymetric distribution. Most speci- 
mens have been collected in less than 40 m and 
some occur to about 90 m; at least some have 
been taken from near estuaries, but no salinity 
information is available. The species are ven- 
omous, as are apparently all scorpionfishes. An 
account of a sting is given by McKay (1964) for 
one species. Our observations on stomach con- 
tents of some species of Inimicus show they feed 
mainly on small fishes. These species lack a gas 
bladder and apparently sit quietly camouflaged 
on the bottom waiting for prey to approach 
(Weber 1913; McKay 1964; Condé 1977). 


ACKNOWLEDGMENTS 


We are grateful to many persons for their co- 
operation in loaning material or assisting during 
visits to their museums: T. Abe, University of 
Tokyo (UT); E. R. Alfred, National Museum of 
Singapore (NMS); A. L. Alverson, University 
of Washington, College of Fisheries (UWCF); 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


M. L. Bauchot, Muséum National d’ Histoire 
Naturelle, Paris (MNHN); M. Boeseman, Rijks- 
museum van Natuurlijke Historie, Leiden 
(RMNH); J. E. Bohlke, Academy of Natural 
Sciences of Philadelphia (ANSP); W. L. Chan, 
Fisheries Research Station, Hong Kong 
(FRSHK); M. M. Dick, Museum of Compara- 
tive Zoology, Harvard University (MCZ); T. H. 
Fraser and M. M. Smith, J. L. B. Smith Institute 
of Ichthyology, Grahamstown, South Africa, 
(RUSI); P. C. Gonzales and A. F. Umali, Na- 
tional Museum, Philippines (NMP); R. J. 
McKay and G. R. Allen, Western Australian 
Museum, Perth (WAM); J. Nielsen, Zoologiske 
Museum, Copenhagen (ZMK); H. Nijssen, 
Zoologisch Museum Amsterdam (ZMA); J. E. 
Randall, Bernice P. Bishop Museum (BPBM); 
V.G. Springer and staff, United States National 
Museum of Natural History (USNM); and A. 
Wheeler, British Museum (Natural History) 
(BMNH). 

A trip to overseas museums by Eschmeyer 
and participation in this study by Hallacher were 
supported by a grant from the National Science 
Foundation (GB-15811). Rama-Rao completed 
his portion of the study while supported by a 
visiting scientist fellowship from the California 
Academy of Sciences and a travel grant from 
the Smithsonian Institution through the U.S. 
Foreign Currency Program. We are grateful to 
the Zoological Survey of India for permitting 
him to participate in this study. 

Illustrations were prepared by Katherine P. 
Smith. M. Giles, K. Lucas, and L. Ullberg as- 
sisted with photography. T. McHugh provided 
Figures 9-10, taken at Steinhart Aquarium. W. 
I. Follett, J. Gordon, T. Iwamoto, D. Kavan- 
augh, J. McCosker, S. Poss, B. Powell, C. 
Ruark, P. Sonoda aided in the study. Lillian 
Dempster assisted with literature work and no- 
menclatural problems and provided comments 
on the manuscript. 


METHODS 


Methods of measuring and counting follow 
Eschmeyer (1969). Pectoral fin length is mea- 
sured from the base of the first ray to the tip of 
the posteriormost ray; in those species of Jn- 
imicus which have the upper one or two rays 
long and filamentous, the filaments are not mea- 
sured as part of the pectoral fin length. In fishes 
of the subfamily Choridactylinae, the last soft 
ray of the dorsal and anal fins is separate, with 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 477 


its own support, and widely spaced from the 
preceding ray. 

Abbreviations of depositories other than those 
listed in the Acknowledgments are: SU—Stan- 
ford University, now housed at the California 
Academy of Sciences (CAS), ZSI—Zoological 
Survey of India, Calcutta, HUJ—Hebrew Uni- 
versity, Jerusalem, QMB—Queensland Mu- 
seum, Brisbane. 


SUBFAMILY CHORIDACTYLINAE KAUP 


Choridactylinae KAupP, 1858:332 (as a subfamily of Triglidae). 
GILL 1888:569 (compiled from Kaup). 

Pelorinae GILL, 1893:135 (as a subfamily of Synanceidae). 
JORDAN AND STARKS 1904:93, 158 (as a subfamily of Scor- 
paenidae). MATSUBARA 1943:413-414 (defined on internal 
features, Inimicus only, as a subfamily of Scorpaenidae). 

Inimicinae GILL, 1905:224 (Pelorinae in synonymy; genera). 
FOWLER 1938a:51 (in key). ESCHMEYER AND RAMA-RAO 
1973:339 (characterization; included genera). 


The subfamily name Choridactylinae appar- 
ently has not been used since it was proposed 
in 1858, except that Gill (1888) quoted Kaup’s 
use of the name. The subfamily name Inimicinae 
has received only limited usage. We do not feel 
the use of Choridactylinae will upset a long-es- 
tablished name or cause confusion. Pelorinae 
cannot be used since Pelor is a junior homonym 
(see Inimicus below). 

SUBFAMILY DIAGNOosIS.—Body and head 
without scales; except lateral line scales buried, 
widely spaced, about 13-15 in number; plus ad- 
ditional buried scales (usually appearing as 
warts or tufted lumps) between lateral line and 
dorsal fin base. Dorsal spines 12—18, dorsal soft 
rays 5-10, total spines plus rays 21-27. Anal 
spines always 2, soft rays 8-13. Pectoral fin rays 
12, lowermost 2 (nimicus) or 3 (Choridactylus) 
entirely free. Pelvic fin with 1 spine and 5 soft 
rays. Most soft fin rays branched. Second sub- 
orbital (third infraorbital) bone becoming wider 
posteriorly, and attaching to preopercle; with 2 
main ridges, one pointing up to third suborbital 
bone, one pointing back; fourth suborbital bone 
absent or fused with third. Vertebrae 26-30. 


Key to the Genera and Species of the 
Subfamily Choridactylinae 


la. Dorsal spines 12-15; 3 lower pectoral rays 
free and detached from remainder of fin 
Choridactylus 2 
1b. Dorsal spines 15—18; 2 lower pectoral rays 
free and detached from remainder of fin 
TInimicus 3 


2a. Dorsal spines 12-14, usually 13; total anal 
fin spines plus soft rays 10-11, usually 
10; pectoral fin without filamentous rays; 
inner pectoral coloration as in Figure la 
Choridactylus multibarbus (Fig. 2a) 
2b. Dorsal spines 13-15, usually 14; total anal 
fin spines plus soft rays 10-12, usually 
11; pectoral fin with first ray ending in 
a long filament; inner pectoral coloration 
ASmineipine be... £1 wee ee eee 
Choridactylus natalensis (Fig. 2b) 
3a. Dorsal spines from fourth to last mostly 
free from interspinous membrane; snout 
usually longer than postorbital distance 
(postorbital distance into snout 0.9-1.7) 


3b. Dorsal spines from fourth to last well con- 
nected by interspinous membrane up to 
midheight of spines; snout shorter than 
postorbital distance (postorbital distance 
into snout 0.7—-0.9) 
4a. Upper two pectoral rays filamentous in 
juveniles and adults; orbits extremely 
elevated, close-set, and broadly joined 
at their bases; interorbital space about 
equal to orbit diameter; inner pectoral 
fin coloration as in Figure Ic _____________- 
Inimicus filamentosus (Fig. 3a) 
4b. Upper two pectoral rays not filamentous 
in specimens over about 50 mm SL; or- 
bits little elevated, widely spaced at their 
bases and joined only by a low ridge of 
bone; interorbital space wide, usually 
greater than 1% times orbit diameter; in- 
ner pectoral fin coloration not as in Fig- 
ure Ic 
Sa. Inner pectoral fin coloration uniform 
gray, without any markings, as in Figure 
lid. 25 See Se Inimicus cuvieri (Fig. 3b) 
5b. Inner pectoral fin coloration not plain col- 
One. i=... — Pee eas Shae weee 2 Fee 6 
6a. (3 choices) Inner pectoral fin with pale 
spots of irregular size on a dark back- 
ground as in Figures te-f = 
Inimicus sinensis (Fig. 3c) 
6b. Inner pectoral fin with two dark areas al- 
ternating with white as in Figures Ig—h 
Inimicus caledonicus (Fig. 4) 
6c. Inner pectoral fin coloration as in Figures 
Iie eae a etd eet 2 A 
seed Inimicus didactylus (Figs. 5, 9-10) 
7a. Inner pectoral fin dusky, with black spots, 
and a transverse white band in middle of 


478 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


Ficure 1. Coloration on inner surface of right pectoral fin in species of Choridactylus and Inimicus. (a) C. multibarbus, 
CAS 15069, 89.5 mm SL; (6) C. natalensis, CAS 31463, 70.3 mm; (c) I. filamentosus, MNHN A998, 178 mm; (d) I. cuvieri, 
CAS 13536, 130mm; (e)/. sinensis, USNM 218534 122 mm; (f)/. sinensis, WAM P4989, 55 mm (note filament in this juvenile); (g)/. 
caledonicus, based on MNHN A2498, 91.2 mm and 96.3 mm; (A) I. caledonicus, SU 20555, 170 mm; (i) I. didactylus, USNM 
168173, 29.4 mm, juvenile; (j) J. didactylus, USNM 136458, 79.8 mm; (k) I. didactylus, USNM 218504, 132 mm; (/) I. didactylus , 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 


fineasminy Hiomne: lime ee ee 

Inimicus brachyrhynchus (Fig. 6) 

7b. Inner pectoral fin coloration not as above, 
dusky to blackish, often with brown or 
black spots or streaks as in Figures In—o 


8a. Third pectoral ray from bottom free for 
about % its length as in Figure In_______- 
Inimicus joubini (Fig. 7a) 
8b. Third pectoral ray from bottom not free, 
except just at tip as in Figure lo__________ 
TInimicus japonicus (Fig. 7b) 


Genus Choridactylus Richardson 


Choridactylus RICHARDSON, 1848:8-10, pl. 2, figs. 1-3 (type- 
species Choridactylus multibarbus Richardson, 1848, by 
monotypy). 

Chorismodactylus: GUNTHER 1860:151 (‘‘unjustified emenda- 
tion’’ for Choridactylus Richardson). 

Choridactylodes GILCHRIST, 1902:101 (type-species Choridac- 
tylodes natalensis Gilchrist, by monotypy; differed from 
Choridactylus in having usually one more dorsal spine and 
a hairlike prolongation of the uppermost pectoral ray). 


Following earlier authors, we recognize two 
species in the genus Choridactylus, and we do 
not feel that the small differences between them 
warrant two genera. Choridactylus natalensis 1s 
restricted to the southeast coast of Africa, and 
C. multibarbus occurs from the Red Sea east to 
China and the Philippines (see Fig. 8). 

DIAGNosIs.—Dorsal XII-XV, 8-10, total 21- 
24. Anal II, 8-10. Pectoral 12, with 3 lower rays 
free. Vertebrae 26-28. Head small, eyes slightly 
elevated, with depressed area at occiput extend- 
ing across nape and behind eyes. Head blunt in 
front, snout not much longer than orbit. Inter- 
orbital space wide, about equal to 1.5 times orbit 
diameter, with a median depressed area. Mouth 
small, slightly oblique. Villiform teeth on jaws, 
none on vomer and palatines. Small tentacles or 
skin flaps present on head and body, especially 
on lower jaw, eye, spinous dorsal and pectoral 
fins, and as tufts on lateral line scales and above 
lateral line. 

SPINATION.—Head shape and spination vir- 
tually the same for both species. Most head 
spines developed as lumps. Lachrymal bone 
with two spines over maxillary, first small, 
pointing out and forward, second very long and 
directed backward, extending past jaw to below 


— 


479 


eye. Two ridges or spines present on lateral face 
of lachrymal bone; suborbital spines poorly de- 
veloped, usually one short ridge or lumps fol- 
lowed by second longer ridge. First two pre- 
opercular spines well developed; first very long, 
with a small supplemental spine usually present 
at its base, second moderate, third through fifth 
small to virtually absent. Nasal spines absent. 
Preocular spines broad, followed by three or 
four small lumps over eye at position of supra- 
ocular and postocular spine. Tympanic spines 
absent. Parietal and nuchal spines present, 
joined; sometimes as three lumps. Pterotic spine 
small. Sphenotic spine absent. Posttemporal 
bone with a long, blunt spine and with a ridge 
or spine present at anteroventral base. Supra- 
cleithral spine small, obscured by skin. Cleithral 
spine present, blunt. Opercular bone with two 
poorly defined ridges. 


Choridactylus multibarbus Richardson 
(Figures la, 2a, 8; Table 1) 


Choridactylus multibarbus RICHARDSON, 1848:8-10, pl. 2, 
figs. 1-3 (original description; type-locality Sea of China). 

Apistus niger: JERDON 1851:141 (listed; Madras, India). 

Choridactylus multibarbis: Day 1875:161—162, pl. 39, fig. 2 
(description; referred Jerdon’s identification of Apistus ni- 
ger to this species; common at Madras). Day 1889:74, fig. 
32 (description; figure from Day 1875). TORTONESE 1934:227 
(description; Massaua, Red Sea). ?BorRopDIN 1932:89-90 
(venom properties and fear by natives; Raitea, Society Is. 
[see Remarks below]). TORTONESE 1937:205 (Red Sea, one 
specimen from Massaua; short description). ? FOWLER 
1938b:290 (listed; Society Is.; from Borodin 1932). 
BLEGVAD 1944:192-193 (listed from Iranian Gulf). HERRE 
AND HERALD 1950:89-90 (Philippines; good figure; two 
from Manila Bay in 13-15 fms). HERRE 1951:466—-488 (brief 
synonymy; range; description of Philippine specimens). 
Munro 1955:250 (description; Gulf of Manaar). SMITH 
1958:176, pl. 7, fig. J (compiled; figure from Richardson 
1848; incorrectly cited Mauritius instead of coast of Arabia 
for Steindachner 1907). SAMUEL 1961:79-83 (‘‘walking”’ 
mechanism; description and osteology of pectoral girdle). 
SMITH AND SMITH 1963:54, pl. 51J (expected at Seychelles; 
figure from Richardson 1848). AHMED AND QURESHI 
1970:205, text-fig. (description; coastal waters and tidal in- 
lets of Pakistan). 

Chorismodactylus multibarbis: GUNTHER 1860:151-152 (in- 
correct subsequent spelling of Choridactylus multibarbus; 
brief description; China, and Madras, India). BOULENGER 
1889:239 (listed; Muscat, east coast of Arabia). STEINDACH- 
NER 1907:163 (listed; east coast of Arabia). 

Chorismodactylus multibarbus: BLEEKER 1873:141 (listed 
from China, ‘‘=Choridactylus multibarbis’’). 


CAS 15159, 110 mm; (m) I. brachyrhynchus, CAS 31424, 109 mm; (n) I. joubini, MNHN 26.398, 158 mm, holotype; (0) /. 
japonicus, based on USNM 86404 (142 mm), CAS 31435 (136 mm), and CAS 15769 (2, 139-191 mm). 


480 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


FIGURE 2. 
(b) C. natalensis, BMNH 1901.9.24.2, 59.8 mm, syntype, Natal, South Africa. 


REMARKS.—Borodin (1932) identified Chori- 
dactylus multibarbus from the Society Islands, 
and Fowler subsequently (1938b) listed this spe- 
cies from there. It is unlikely that C. multibarbus 
occurs there as the genus is not otherwise 
known from Oceania. The fear by the natives 
and capture by spearing suggests a stonefish 
(Synanceia). 


(a) Choridactylus multibarbus, BPBM uncat., 83 mm SL, Madras, India (photo by J. Randall of a fresh specimen); 


Gunther (1860) seems to have been the first to 
misspell the genus and species name (as .Chor- 
ismodactylus multibarbis), and most subsequent 
workers have wrongly followed him. The origi- 
nal correct spelling is Choridactylus multibar- 
bus. 


MATERIAL EXAMINED.—Persian Gulf: BPBM 21081 (1, 87). 
Southeast coast of India: ANSP 77541 (1, 57.8). MNHN 02- 


481 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 


ec Il — a a cl ce I L8°EC = a t ve 6 = = 
Os Ol a ra = = I I 00°FC = = = Cc — a ae 
00°CI a = x Cc = = 00°SC = 7 c = — a = 
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st el a t ¢ a = = CC'S a G iL = a = = 
00'rI c CE c os =< ms ELRSG = OC 9 a = = => 
80°F E Ip (e = - = 88°SC i bP fe = = = = 
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00'TT = — = I 9 I OS TC =a = . I ie £ I 
90°01 = = = = I I 00°TC = 3 a = I rl I 
x Sl rl €l cl Il Ol us LC OG SC PC tC CC I7 
skey pue soulds jeuy [P10], sAey pure soulds [esiog [R10], 
169 ae 2 iS tt L I 86°91 v Lt iS = oz a ae 
O00'L = x = G + = 00°LI = c = = — — = 
00°8 — ar G = = 7 00°LI = G = = — 1 = 
978 = el €C € — a 00°91 = c Sa Cc — =; = 
CC'8 i Cc L — = = 00°LI ra 6 a ea ae ae = 
crs =" cl tl I =r ary ce Li 6 Ll a = a =z ima 
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skey IOs [esi0og 


“snoimiuy GNV snjAJIDpiuoy) VYANAD AHL JO SaAINddS NI SLNNOD AVY NIY IVNY GNV Tvsuod “| ATaV 


souids jesiog 


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SNYIUAYAKYIDAG * 
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LA1AND *] 
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snqanqgujniu *) 


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snouodnl 7 
ruiqnol *] 
SNYIUAYAKYIDAG “T 
SN}XIODPIP “] 
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snsojUuauMDpy *] 
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482 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


196 (1, 59.6). SU 14665 (4, 53.9-81.6). CAS 31462 (1, 71.2). 
CAS 33951 (2, 58.0-71.5). 

Gulf of Thailand: CAS 15066 (1, 76.1). CAS 15067 (1, 73.3). 
CAS 15068 (1, 84.3). CAS 15069 (1, 89.5). CAS 15070 (1, 96.0). 
CAS 15071 (1, 76.2, cleared and stained). CAS 15072 (1, 98.5). 

China: BMNH uncat. (51.0, dried, holotype of Choridac- 
tylus multibarbus), ‘‘Samarang,’’ J. Richardson. 

Philippines: UWCF 10164 (1, 51.4). 


DIAGNOsIS.—First pectoral ray not filamen- 
tous in adults; inside of pectoral fin with about 
5 oblong white areas, surrounded by black or 
brown pigment and usually with white specks in 
axil (Fig. la). 

DESCRIPTION.—Dorsal fin with 12—14 (usually 
13) spines and 8—9 (usually 9) soft rays, total 21— 
23 (usually 22). Anal fin with 2 spines and 8-9 
(usually 8) soft rays. Pectoral fin with 12 rays, 
lower 3 rays free, rays 2 through 6 or 7 usually 
branched in adults. Gill rakers on outside of first 
arch total 9-11, 2-3 on upper arch, 7—9 on lower 
arch. Buried lateral line scales about 13-15, in- 
cluding one on caudal fin. Vertebrae 26 (9 spec- 
imens). Head spines and head shape as for the 
genus. 

Coloration in life as given by Day (1875:161; 
1889:74) for a specimen from India: ‘“‘brownish, 
with a yellow shoulder mark, and two or three 
vertical orange bands: base of ventral and anal 
with fine white spots. Fins blackish brown, with 
a light band between the fourth and sixth dorsal 
spines: margins of pectorals orange: caudal with 
a black band at its base, and another in its last 
third having a light edge: outer two-thirds of anal 
blackish: free rays black, with white in the mid- 
dle:* 

General body shape and coloration in preser- 
vative as in Figure 2a. Color variable; head and 
body mostly brown or dark gray, with pale 
areas; sometimes irregular dark spots on sides. 
Dorsal fin usually paler at front, at about spines 
3-5, and at front of soft dorsal fin; pale areas 
usually extending on to body. Caudal fin pale at 
middle (sometimes crossed by narrow vertical 
wavy lines), broadly dark on posterior half, with 
tips of rays pale. Anal and pelvic fins dark, with 
many small pale spots at base and on adjacent 
parts of body. Inner surface of pectoral fin as in 
Figure la; outer surface of pectoral fin mostly 
dark, with paler areas showing through faintly 
from inner side of fin. 

Measurements for three specimens (51.4—75.4 
mm SL) in percent SL: head 35-36, snout 11- 
14, orbit 9-11, interorbital width 11-12, post- 
orbital 15-17, pectoral 37-39. 


COMPARISONS.—The two species of Chori- 
dactylus can be separated from all other scor- 
paenids by the combination of three free pec- 
toral fin rays and no scales on the body. 
Choridactylus multibarbus differs from C. na- 
talensis by lacking the elongated filamentous 
first pectoral ray (although small specimens of 
multibarbus may have a slight elongation of this 
ray), having one less vertebra (26 versus 27), 
and in having a different coloration on the inside 
of the pectoral fins (Fig. la versus 1b). Chori- 
dactylus multibarbus averages about one fewer 
dorsal spines, one more dorsal soft ray, and one 
less anal soft ray than C. natalensis. 

DISTRIBUTION.—Choridactylus multibarbus 
is known from the Red Sea, Persian Gulf, Gulf 
of Oman, Pakistan, India, Gulf of Thailand, 
China, and the Philippines. It is known from 
sand or mud bottoms in depths from near shore 
to 40 m. 


Choridactylus natalensis (Gilchrist) 
(Figures 1b, 2b, 8; Table 1) 


Choridactylodes natalensis GILCHRIST, 1902:101-103, pl. V, 
figs. 1 and 2 (original description; as type of a new genus; 
two specimens from 2!4 miles [4 km] off Umhlanga River 
mouth, in 22-26 fms [40-48 m], bottom of fine sand, 25 Mar. 
1901; and five specimens from off Cape Natal Lighthouse, 
in 22 fms [40 m], bottom of fine sand). GILCHRIST AND 
THOMPSON 1917:410 (listed, Natal). VoN BONDE 1924:3, 30 
(specimens from SE coast of Africa, PICKLE stations 169, 
190, and 193, depths 27-29 fms [49—53m]). FOWLER 
1925:254 (bref description; Durban Bay [ANSP 86316]). 
FOWLER 1934:485 (listed from Durban and Natal [ANSP 
87807, 86316, 88020]). SMITH 1949:375, pl. 83 (brief descrip- 
tion; Natal; color fig.). 

Choridactylus natalensis: BARNARD 1927:922, pl. XXXIII 
(description; synonymy; figure from Gilchrist 1902). BAR- 
NARD 1947:199, pl. 24, fig. 2 (compiled). SMITH 1958:176— 
177, pl. 8, fig. B (description; coloration; two specimens 
from 20 fms [37 m], off Natal; good photograph). SMITH 
AND SMITH 1963:54, pl. 52, fig. B (expected at Seychelles; 
figure from Smith 1958). 


MATERIAL EXAMINED.—AII specimens from the SE coast 
of Africa: BMNH 1901.9.24.2 (60.0, syntype of C. natalensis) 
3 miles [4.8 km] off Natal coast, collected by J. D. F. Gilchrist. 
ANSP 86316 (1, 108). ANSP 87807 (1, 103). ANSP 88020 (1, 
63.5). RUSI 2000 (2, 65.5-67.0). RUSI 2001 (1, 80.2). CAS 
31463 (1, 70.3). 


DIAGNOSIS.—First pectoral ray terminating in 
a long filament, extending well beyond margin 
of fin; inside of pectoral fin with dark stripes 
(over rays) on a pale background (Fig. 1b). 

DESCRIPTION.—Dorsal fin with 13—15 (usually 
14) spines and 8-10 (usually 8 or 9) soft rays, 
total 21-24 (usually 22-23). Anal fin with 2 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 


spines and 8-10 (usually 9) soft rays. Pectoral 
fin with 12 rays, lower 3 rays free, rays 2-5 
branched in available material. Gill rakers on 
outside of first arch total 8-10, 1-3 on upper 
arch, 6-8 on lower arch. Buried lateral line 
scales 13-14, including one on caudal fin. Ver- 
tebrae 27 (3 specimens) or 28 (1). Head spines 
and head shape as for the genus. 

Coloration in life as given by Smith (1958:177): 
‘*Colour variable, brilliant, mottled, spotted and 
marbled in brown, pink and yellow. Front of 
body rather darker, several vague bars on hinder 
part extending onto dorsal and anal. Base and 
distal 3rd of caudal dark, mid posterior and hind 
margin pink. Lower free pectoral rays annulate 
pink.” 

General body shape and coloration in preser- 
vative as in Figure 2b. Color variable; body and 
head mostly brown, mottled, sometimes with 
darker brown spots on side of body. Dorsal fin 
brown, paler at level of spines 4-6, 11-12, and 
on anterior half of soft dorsal fin. Caudal fin pale 
in middle, a broad dark band on posterior half, 
with tips of rays or distal edge of fin pale. Anal 
fin brown, usually mottled with lighter pigment. 
Pelvic fins mostly dark brown, often with minute 
pale dots at base and on adjacent part of body. 
Inner surface of pectoral fin with dark bars over- 
lying the rays (Fig. 1b); outer surface of pectoral 
fin brown or tan, mottled with darker and lighter 
areas. 

Measurements for four specimens (65.5—103 
mm SL) in percent SL: head 33-35, snout 10- 
12, orbit 7-10, interorbital width 8-10, postor- 
bital 15-17, pectoral fin 31-37. 

COMPARISONS.—See account of C. multibar- 
bus (p. 482). 

DISTRIBUTION.—Choridactylus natalensis 1s 
known only from Durban Bay, South Africa, 
north to Mozambique, in depths from about 35 
to 75 m on sandy bottom. 


Genus Inimicus Jordan and Starks 


Pelor CUVIER, 1829:168 (type-species Scorpaena didactyla 
Pallas, 1769, by original designation [footnote 6 on p. 168 
as, ‘‘Pel. obscurum, Nob., ou Scorpaena didactyla, Pall., 
Spic. Zool, ... . ")))s 

Inimicus JORDAN AND STARKS, 1904:158 (type-species Pelor 
Japonicum Cuvier in Cuvier and Valenciennes 1829, by 
original designation; wrongly thought type-species of Pelor 
was filamentosum). CONDE 1977:15-16 (species of genus). 

Simopias GILL, 1905:224 (a replacement name for Pelor; type- 
species Scorpaena didactyla Pallas by reason that Simopias 
was proposed as a replacement name for Pelor but Gill 


483 


wrongly thought filamentosum was the type-species of Pe- 
lor; Simopias proposed as a subgenus of Inimicus). 

Chorismopelor CHEVEY, 1927:222 (type-species Chorismope- 
lor joubini Chevey, by original designation; comparison 
with other genera). 


REMARKS.—The generic name Pelor Cuvier 
is preoccupied by Pelor Dejean, 1828 [based on 
unpublished table of Bonelli; see Mroczkowski 
1977:61], a genus of carabid beetles. Jordan and 
Starks (1904:158) proposed the generic name 
Inimicus to accommodate forms like Pelor ja- 
ponicum that do not possess filamentous upper 
pectoral rays. They wrongly assumed the type- 
species of Pelor was filamentosum, the only spe- 
cies then known to have the upper pectoral rays 
filamentous; they did not propose the name Jn- 
imicus aS a replacement name for Pelor, con- 
trary to what Jordan (1917:127) would lead one 
to believe. The type-species of Pelor was estab- 
lished by Cuvier when he first proposed the 
name (1829:168) as Scorpaena didactyla (which 
Cuvier preferred to list under his own new name 
obscurum). Gill (1905:224) also wrongly as- 
sumed the type-species of Pelor, and hence his 
replacement generic name Simopias, was fila- 
mentosum as indicated by the last sentence un- 
der his treatment of /nimicus; and he therefore 
confused his subgeneric designations. His con- 
fusion over the type-species does not obviate his 
replacement of Pelor with Simopias. His state- 
ments, ““The name Pel/or can not be retained for 
this genus inasmuch as it had been appropriated, 
in 1813 by Bonelli. . . , > and ‘‘The name [Sim- 
opias| here proposed as a substitute... .’’ are 
a definite indication that he was replacing the 
preoccupied Pelor. Inimicus, by reason of prior- 
ity, takes precedence over Simopias. 

Chorismopelor was proposed for the species 
jJoubini, which has the third pectoral ray from 
the bottom free for part of its length, unlike the 
other species that have the third ray from the 
bottom fully connected by membrane to the ray 
above it. We do not feel this species warrants 
a separate genus (see Species Characters be- 
low). 

We recognize eight species of Inimicus. The 
genus is mostly coastal in distribution, occurring 
from the western Indian Ocean to Oceania (Fig. 
8). 

D1AGNosis.—Dorsal XV-XVIII, 5-9, total 
23-27; first three dorsal spines fully joined by 
membrane, separated from rest of fin but con- 
nected by membrane to fourth spine basally; re- 


484 


maining spines long and sharp, of about uniform 
height (except at beginning and end), nearly free 
of interspinous membranes or connected to 
about midheight. Anal II, 8-13. Pectoral 12, 
lower two rays free, third from bottom free for 
up to one-third its length in one species. Ver- 
tebrae 27-30. Eyes elevated, partially or almost 
fully joined by a bony ridge, with a shallow to 
deep occipital depression behind. Large, round 
to oblong depressed area in front of each eye, 
separated by an elevated median ridge running 
from base of eyes to snout. Mouth large, up- 
turned. Small canine teeth on jaws and vomer, 
no teeth on palatines. Small tentacles or skin 
flaps present on head and body, especially on 
lower jaw, eye, spinous dorsal and pectoral fins, 
and as tufts on lateral line scales and above lat- 
eral line. 

SPINATION.—Head spines moderately devel- 
oped and covered by thick skin and often fringed 
with cirri. Lachrymal bone with two blunt spines 
over maxillary, first small and pointing forward 
and down, second larger and pointing down and 
back. Suborbital ridge with 4 spinous lumps, 
first on lateral face of lachrymal bone, second 
on first suborbital (second infraorbital) bone, 
third and fourth as a pair (one above the other) 
below eye. Preocular, supraocular, and postoc- 
ular spines usually poorly defined. Nasal spine 
small or absent. Preopercle with supplemental 
and first two preopercular spines well devel- 
oped, third through fifth poorly developed or 
virtually absent. Small spine below posterior 
margin of orbit sometimes present. Parietal and 
nuchal spines small, joined, sometimes as a 
rough ridge. Tympanic spine absent. Pterotic 
spine low and blunt, just behind orbit. Upper 
and lower posttemporal spines well developed. 
Supracleithral spine shelflike. Cleithral spine 
virtually absent. Opercle with one broad spine. 

SPECIES CHARACTERS.—Meristic characters 
which differ among the species include dorsal 
and anal fin rays and vertebrae. Number of pec- 
toral rays and pelvic rays is the same in all spe- 
cies. Measurements and ratios of body parts can 
be used in separating some species, the most 
useful of which are the length of dorsal spines 
in relation to the height of their interspinous 
membranes; interorbital width in relation to or- 
bit diameter; and snout length in relation to post- 
orbital distance. Juveniles apparently have the 
upper one or two pectoral rays elongated as fil- 
aments, but these are retained only in the adults 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


of 1. filamentosus. Coloration on the inner sur- 
face of the pectoral fin is very useful in species 
identification. 

Three species groups are recognizable. Jnim- 
icus filamentosus is distinct in retaining in adults 
the condition of the upper pectoral rays elon- 
gated into filaments and in having strongly ele- 
vated, close-set, and broadly joined orbits. Jn- 
imicus caledonicus, I. cuvieri, I. didactylus, and 
I. sinensis exhibit similar characters by having 
the snout longer than or about equal to the post- 
orbital distance; the orbits elevated but well sep- 
arated and joined only by a low ridge of bone 
(more pronounced in J. didactylus); and the in- 
terspinous membranes from the fourth to the 
penultimate dorsal spines very low, connecting 
each spine to the base of its succeeding spine; 
and high counts of total dorsal and especially 
anal rays. Inimicus brachyrhynchus, I. japoni- 
cus, and I. joubini resemble each other by hav- 
ing the snout shorter than the postorbital dis- 
tance; the orbits elevated but widely spaced, and 
joined only by a low ridge of bone, as in the 
previous group; the interspinous membranes be- 
tween the dorsal spines extending up to about 
the midheight of the spines; and a low anal fin 
ray count. Inimicus joubini differs from the oth- 
er two members of this third group by having 
the third pectoral ray from the bottom free dis- 
tally for part of its length. 


Inimicus filamentosus (Cuvier) 
(Figures Ic, 3a, 8; Table 1) 


Pelor filamentosum CuvieR, 1829:428-434, pl. 94 (original 
description; type-locality Mauritius). LEsson 1831:206, pl. 
21, fig. 1 (Mauritius). CUVIER 1834:285 (Maunitius; Squilla 
as food). CUvIER 1836:pl. 25, fig. 2 (figure only). CUVIER 
1837: Pisces, pl. 16, fig. 1 (figure poor). KAUP 1858:334 (list- 
ed). GUNTHER 1860:149-150 (brief description; one speci- 
men from Mauritius). BLEEKER 1874b:87 (listed; Borbonia 
[Réunion], Madagascar, and Mauritius). PETERS 1876a:439 
(listed; Mauritius). SAUVAGE 1878:147 (mentioned in type 
description of Pelor caledonicum). SAUVAGE 1891:517 (list- 
ed; Madagascar). JORDAN AND STARKS 1904:158 (men- 
tioned). BLANC AND HUREAU 1968:27 (type at MNHN). 

Pelor filamentosus: GUICHENOT 1863:24 (listed; Réunion I.). 
GILL 1905:224 (as type of Simopias). 

Pelor didactylum: REGAN 1908:237 (misidentified; listed; Mal- 
dives, in 30 fms [55 m]). 

Inimicus filamentosus: FOWLER 1938a:86 (briefly compared to 
I. bifilis). SM1TH 1958:176, pl. 8c (synonymy; description; 
one specimen from Zanzibar; range). McKay 1964:10-11 
(compared to /. bifilis). TORTONESE 1968:25 (Red Sea; seen 
in Eilat Aquarium; compiled range). CONDE 1977:15-18, 
figs. 1-4 (description; excellent color photographs; behavior 
and aquarium observations). 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 


Ficure 3. (a) Inimicus filamentosus, HUJ 5261, 72 mm SL, Eilat, Red Sea; (b) J. cuvieri, NMS 1359, 113 mm, South China 
Sea; (c) J. sinensis, FRSHK uncat., 88.2 mm, Hong Kong. 


MATERIAL EXAMINED.—Mauritius: MNHN 6714 (about 
148, holotype of Pelor filamentosum), Ile de France [Mauri- 
tius], collected by Lesson and Garnot, Duperrey Expedition. 

Red Sea: BPBM 18283 (1, 171). HUJ—F-5261 (1, 72). 

Zanzibar: RUSI 1999 (1, 122). 

Malagasy [Madagascar]: MNHN A998 (1, 178). 

Reunion I: MNHN 1966-854 (1, 165). 

DIAGNOSIS.—Two uppermost pectoral rays 
elongated as filaments in specimens of all sizes; 
filament on second ray longer than first. Orbits 
extremely elevated, close-set, and broadly 

joined at their bases. Interorbital distance about 
equal to orbit diameter. A deep, oblong pit be- 
low and in front of eye. Occiput depressed, 
forming a deep saddle behind orbits. Snout lon- 
ger than postorbital distance (postorbital into 
snout |.2—1.3). Interspinous membrane from 
fourth dorsal spine rearwards less than one-half 
spine height. Inner pectoral fin coloration (Fig. 
lc) diagnostic. 

DESCRIPTION.—Dorsal fin with 15—16 (usually 
15) spines and 7-8 (usually 8) soft rays, total 23. 
Anal fin with 2 spines and 9-10 (usually 10) soft 
rays. Gill rakers on outside of the first arch total 
8-11, 3 on upper arch, 5-8 on lower arch. Ver- 
tebrae 27 (4 specimens). Head spines and gen- 
eral features as for the genus. 

Color in life, shown in Condé (1977:15-18), 
mostly brown and yellow. Inner surface of pec- 
toral fin mostly brilliant yellow, with black 
patches at base, brown spots subterminally, and 
black distally. 

General body shape and coloration of pre- 
served specimens as in Figure 3a. Body and 
head brown, mottled with pale areas. Large pale 
bars extend down on sides from dorsal fin in 
some specimens. Area before eyes and in occi- 
put with large circular pale spots. Dorsal spines 
tipped with black or dark brown. Soft dorsal, 
caudal, and anal fins darker distally. Pelvic fins 
dark brown. Outer surface of pectoral fin brown 
to black surrounding a large pale area in middle. 
Inner surface of pectoral fin distinctively marked 
(Fig. lc); upper three to five rays with black or 
dark brown patch just above base, sometimes as 
two or three roundish dark spots; distal third of 
fin with brown or black spots, grading into solid 
black or brown at edge. 

Measurements for four specimens (72.2—178 
mm SL) in percent SL: head 36-38, snout 18- 
19, orbit 6-7, interorbital width 5—9, postorbital 
15-16, pectoral fin 41-50. 

DISTRIBUTION.—Inimicus filamentosus is 
known from the Red Sea and the western Indian 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


Ocean; specific localities include Eilat, Gulf of 
Aqaba, Zanzibar, Malagasy Republic (Mada- 
gascar), Réunion I., Mauritius, and Maldives. 
Little is known of its depth distribution; one rec- 
ord is in 55 m, but the species probably occurs 
to nearshore depths, presumably on sand bot- 
tom. 


Inimicus cuvieri (Gray) 

(Figures 1d, 3b, 8; Table 1) 

Pelor cuvieri GRAY, 1835:pl. 90, fig. 2 (drawing based on a 
specimen from Singapore). BLEEKER 1852a:252—253 (de- 
scription; Priaman, Sumatra). BLEEKER 1854a:57 (listed; 
Priaman, Sumatra). GUNTHER 1860:150 (in part, sinense 
wrongly included; brief description). BLEEKER 1861:49 (list- 
ed; Singapore). BLEEKER 1873:141 (listed; China; possibly 
synonymous with P. sinense). BLEEKER 1874a:4, 5, 8-10, 
pl. 2, fig. 3 and pl. 3, fig. 1 (description; good figures; Su- 
matra). 

Inimicus cuvieri: MCKay 1964:10-11 (brief reference to pre- 
vious literature; suggestion of possible filamentous pectoral 
rays in juvenile /. cuvieri). 


MATERIAL EXAMINED.—Singapore: BMNH uncat. (about 
172, holotype of P. cuvieri), see Remarks below. SU 39471 
(1, 53.5). 

South China Sea: NMS 1358 (1, 93). NMS 1359 (1, 113). 

Indonesia: NMS 738 (1, 113), Sarawak. 

Gulf of Thailand: USNM 10334 (1, 195). The following were 
collected by the George Vanderbilt Foundation in 1960-61, 
most between 11°50’N and 99° to 100°E: CAS 13531 through 
CAS 13559 and CAS 13563 and CAS 31182, 40 specimens, 94— 
194 mm SL. The following were collected by F. Steiner: CAS 
13560 (1, 92), CAS 13561 (3, 101-137), CAS 13562 (6, 88-153), 
and CAS 34299 (1, 109). 


REMARKS.—The labels with the British Mu- 
seum specimen indicate that it is the type and 
was collected by Hardwicke in India. This spec- 
imen corresponds to bottle ‘‘a’’ in Giinther’s 
1860 ‘Catalogue.’ The type-locality given by 
Gray (1835) for the illustrated specimen was Sin- 
gapore as published in ‘‘Illustrations of Indian 
Zoology.’’ Perhaps the locality India on the bot- 
tle label resulted from the title of the publication. 
To our knowledge, /. cuvieri does not occur off 
India. 

DIAGNOosIs.—No filamentous pectoral fin rays 
in adults. Orbits only slightly elevated. Snout 
length usually equal to or longer than postorbital 
distance (postorbital distance into snout 0.9- 
1.3). Interspinous membrane from fourth dorsal 
spine rearwards less than one-fourth spine 
height. Inner surface of pectoral fin (Fig. 1d) 
without distinctive pattern, mostly solid brown, 
without spots. 

DESCRIPTION.—Dorsal fin with 17-18 spines 
and 8-9 soft rays, total 25-27 (usually 26). Anal 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 487 


fin with 2 spines and 11—13 (usually 12) soft rays. 
Gill rakers on outside of first arch total 8-11, 2- 
3 on upper arch, 6-8 on lower arch. Vertebrae 
29 (10 specimens), or 30 (1). Head spines and 
general body features as for the genus. Skin on 
occiput and before eyes spongy, more so than 
in other species, thereby minimizing the appar- 
ent depth of occiput and pits before eye. Up- 
permost one or two pectoral rays elongated in 
juveniles, but not in specimens longer than 
about 55 mm SL. 

General body shape and coloration of pre- 
served specimens as in Figure 3b. Head and 
body mostly brown or gray-brown above, paler 
below. Some specimens with much of body pale, 
with dark brown on and just behind head, at 
middle and end of spinous dorsal fin, and below 
end of soft dorsal fin. Head usually with minute 
dark spots. Pale spot or spots present before 
eyes in some specimens. Soft dorsal fin usually 
pale anteriorly. Caudal fin mostly dark brown or 
black, often pale at base and at center, alternat- 
ing with dark. Anal fin and pelvic fins darker 
distally. Inner surface of pectoral fin mostly dark 
brown to black, mostly plain colored, without 
round spots or stripes, although tiny pinpoint- 
sized pale spots sometimes present. Outer sur- 
face of pectoral fin brownish to blackish, some- 
times with tiny brown spots and large vague pale 
areas. 

Measurements for thirteen specimens (92.4— 
194 mm SL) in percent SL: head 31-40, snout 
14-16, orbit 4-6, interorbital width 8-10, post- 
orbital 12-16, pectoral fin 33-43. 

COMPARISONS.—lnimicus cuvieri and I. si- 
nensis are very similar in all features except that 
I. sinensis has large yellow spots (pale in pre- 
servative) on the inner side of the pectoral fin, 
while J. cuvieri lacks spots and has the inner 
surface of the pectoral fin more or less uniformly 
dusky. 

DISTRIBUTION.—/nimicus cuvieri is confined 
to the area from the Gulf of Thailand to Sara- 
wak, Sumatra, and Singapore. It is evidently 
very common in the Gulf of Thailand on sandy 
or mud-sand bottom. Depths of capture for the 
George Vanderbilt Foundation specimens were 
from 18 to 50 m, with one reportedly from 1 m. 


Inimicus sinensis (Valenciennes) 
(Figures le-f, 3c, 8; Table 1) 


Pelor sinense VALENCIENNES in CUVIER AND VALEN- 
CIENNES, 1833:468—469 (original description; type-locality 


Canton, China [holotype MNHN 6374]). RICHARDSON 
1846:212 (listed; Canton, China). GUNTHER 1860:150 (in 
part; listed as possible synonym of P. cuvieri; China). 
BLEEKER 1873:141 (listed as possible synonym of P. cuvieri; 
Canton, China). SAUVAGE 1873:53, pl. 7, fig. 2 (description 
and figure based on type; China). 

Pelor cuvieri (not of Gray): RICHARDSON 1844:72-73, pl. 
XXXIX, fig. 4 (description; good figure; China); 1846:212 
(brief synonymy; Canton, China). 

Inimicus cuvieri (not of Gray): HERRE AND HERALD 1950:355 
(listed; Philippines). FOWLER 1931:305 (Hong Kong); 
1938d:30 (listed; Hong Kong). 

Inimicus cirrhosus MCKAY, 1964:8-12, fig. 1 (original descrip- 
tion; type-locality Shark Bay, Western Australia). 

Inimicus sinense: RAMA-RAO AND BADRUDEEN 1973:418—421, 
fig. 1 (synonymy; description; Ceylon and SE coast of In- 
dia). 


MATERIAL EXAMINED.—Ceylon: USNM 218534 (1, 122). 

Java: CAS 36066 (2, 60-61). MCZ 1097 (1, 140), ZMK P- 
791105 (1, 55.8). 

Viet Nam: CAS 31423 (1, 83.8). 

Hong Kong: CAS 31422 (1, 144). ANSP 76624 (1, 136). 
ANSP 76707 (1, 153). FRSHK uncat. (1, 88.2). FRSHK uncat. 
(1, 130). SU 60861 (1, 216). SU 61097 (1, 142). 

Taiwan: CAS 15614 (1, 141). 

China: MNHN 6374 (124, holotype of P. sinense). CAS 
27662 (1, 115). 

Philippine Is.: ANSP 48776 (1, 134). BPBM 21083 (1, 114). 
CAS 33339 (1, 61). NMP 1457 (1, 78.6). NMP 2346 (1, 129). 
NMP 8217 (2, 119-120). SU 39135 (1, 108). USNM 99786 (1, 
118). USNM 218532 (1, 98.9). USNM 218533 (1, 136). UWCF 
7215 (3, 76.3—138). 

Australia: WAM P.4981 (1, 150), P.4989 (1, 55.0), and P.4994 
(1, 118), all paratypes of J. cirrhosus, Shark Bay, Western 
Australia. 


DIAGNosIs.—No filamentous pectoral fin rays 
in adults. Orbits only slightly elevated. Snout 
length equal to or longer than postorbital dis- 
tance (postorbital distance into snout 1.0-1.5). 
Interspinous membrane from fourth dorsal spine 
rearwards less than one-fourth spine height. 
Coloration of inner surface of pectoral fin vari- 
able (Figs. le-f), but always with some large 
pale spots. 

DESCRIPTION.—Dorsal fin with 17-18 spines 
and 7-9 (usually 8—9) soft rays, total 25-26. Anal 
fin with 2 spines and 11-13 (usually 12) soft rays. 
Gill rakers on outside of first arch total 7-10, 2- 
3 on upper arch, 5—8 on lower arch. Vertebrae 
28 (1 specimen) or 29 (5). Head spines and gen- 
eral body features as for the genus. Upper pec- 
toral fin rays ending in filaments in juveniles 
(Fig. If) but not filamentous in adults. 

Color in life, according to McKay (1963:9) for 
Australian specimens: ‘‘Pectoral fin dark brown, 
almost black with indistinct pale cross-bars and 
variable blotches on outer surface. Inner surface 
dark brown to blackish, with 20 to 35 white (in 


488 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


\ iz: 


a 
> i. Say, 


Fay ih 


FiGurRE 4. Inimicus caledonicus, ZSI 1749, 101 mm SL (from Day 1875, pl. xxxix, fig. 1). 


life bright yellow) spots of indefinite shape. In 
larger examples the yellow spots are more dif- 
fuse and frequently in the form of ocelli.’’ Rama- 
Rao and Badrudeen (1973:420) provide the fol- 
lowing for a specimen from India: ‘‘Colour in 
life: Body blackish with orange blotches; ocelli 
on the inner surface of pectoral fin and spots on 
caudal fin with yellowish tinge in middle and 
reddish orange around; pelvic fin blackish.” 
General body shape and coloration of pre- 
served specimens as in Figure 3c. Body and 
head dark brown to tan or gray, paler below. 
Sometimes body almost uniform in color, but 
usually mottled with paler areas; some dark 
grayish specimens are strongly marked with 
white patches. Head often with minute dark 
spots, large pale spot before each eye in some 
specimens; body often with irregular small dark 
spots on a pale background or with pale areas 
if background is dark. Soft dorsal fin usually pal- 
er anteriorly. Caudal fin usually with a narrow 
pale area at base and a broad pale area at middle 
of fin, often as band of vague white spots, oth- 
erwise fin dark. Pelvic fin dark distally. Outer 
surface of pectoral fin variable, often reflecting 
white spots of inside. Inner surface of pectoral 
fin mostly dark with large white, somewhat ir- 
regular spots as in Figures le, f; sometimes 
small dark spots also present; large white spots 
variable in shape and in number, from 2 or 3 to 
more than 35; in largest specimens, white spots 


may have a dark center; spots sometimes rather 
faint. 

Measurements for twenty specimens (55.0— 
216 mm SL) in percent SL: head 33-40, snout 
14-18, orbit 4-6, interorbital width 7-12, post- 
orbital 13-17, pectoral fin 33-43. 

COMPARISONS.—See [nimicus cuvieri (p. 487). 

DISTRIBUTION.—Jnimicus sinensis 1s known 
from India, Ceylon, Java, Viet Nam, Hong 
Kong, China, Philippines, and Western Austra- 
lia. Depths of capture range from 5 to about 90 
m, on sand bottom. 


Inimicus caledonicus (Sauvage) 
(Figures lg & h, 4, 8; Table 1) 


Pelor didactylum (not of Pallas): DAY 1875:160-161, pl. 39, 
fig. | (description; Andaman Is.). DAy 1889:72-73, fig. 31 
(compiled from Day 1875). 

Pelor caledonicum SAUVAGE, 1878:147-148, pl. II, fig. 6 (orig- 
inal description; type-locality New Caledonia). FOWLER 
1928:299 (compiled from Sauvage 1878). 

Pelor barbatus DE Vis, 1885:547 (original description; type 
locality Cape York, Australia). MCCULLOCH 1916:96—97, 
pl. 58 (good description; Queensland localities; 9-33 fms 
[16-60 m]). 

Inimicus barbatus: MCKAY 1964:10 (body proportions and in- 
ner pectoral fin coloration; compared with J. didactylus). 
MARSHALL 1965:431-—432, pl. 58, fig. 418 (short synonymy; 
description; distribution). 

Inimicus caledonicus: McKay 1964:10 (counts; compiled). 


REMARKS.—We believe that the single speci- 
men in MNHN 1518 is the holotype of P. cale- 
donicum. It was not listed as a type by Blanc 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 489 


and Hureau (1968), and it carried the specific 
name sinensis when it was made available to us. 
It agrees in locality, size, and in all features with 
the figure and description as given by Sauvage 
(1878) except that the membranes between the 
dorsal spines are illustrated as joining the spines 
for their entire length, a condition not found in 
any species of Inimicus. 

The type of J. barbatus (QMB 112/374) and a 
second Australian specimen (SU 20555) closely 
resemble our specimens from New Caledonia, 
and we feel that barbatus is a synonym of ca- 
ledonicus. We also refer the specimens (ZSI 
1749-1750) reported from the Andaman Islands 
(Day 1875) and a specimen from the Nicobar 
Islands (ZSI 440/2) to I. caledonicus. 


MATERIAL EXAMINED.—New Caledonia: MNHN 1518 (123, 
holotype of P. caledonicum), received from ‘*Musee des Col- 
onies.”” MNHN A2498 (2, 91.2—96.3). MNHN A2833 (1, about 
130, dried). MNHN A4877 (2, 86.8—-90.3). 

Australia: QMB 112/374 (144, holotype of P. barbatus), 
Queensland, Cape York. ANSP 98707 (1, 110), Queensland. 
SU 20555 (1, 170), Queensland. 

Andaman Is.: ZSI F1749-1750 (2, 55.0—101) purchased from 
F. Day [specimen 1749 figured in Day 1875, pl. 39, fig. 1]. 

Nicobar Is.: ZSI F440/2 (1, 122). 


DIAGNOsIs.—No filamentous pectoral rays in 
adults. Orbits only slightly elevated. Snout 
length greater than postorbital distance (post- 
orbital distance into snout 1.2—1.4). Interspinous 
membranes from fourth spine rearwards low, 
extending up to about one-fifth of spine height. 
Inner surface of pectoral fin usually with two 
broad dark areas (Fig. Ig, h). 

DESCRIPTION.—Dorsal fin with 17 spines and 
8-9 (usually 8) soft rays, total 25-26. Anal fin 
with 2 spines and 11-12 soft rays. Gill rakers on 
outside of first arch total 10-12, 3 on upper arch, 
7-9 on lower arch. Vertebrae 28 (2 specimens). 
Head spines and general features as for the ge- 
nus. 

Color in life, according to Day (1875:161) for 
specimens from the Andaman Islands: ‘*brown- 
ish-grey becoming dirty white beneath, with fine 
spots over the body and head. Dorsal coloured 
as the body, a dark band passes down the last 
few spines on to the body, and another over the 
last few rays takes the same course: caudal yel- 
low with a dark vertical band across its base, 
and another in its last third: outer edge of anal 
dark coloured.’’ De Vis (1885:547) provides the 
following for a specimen from Australia: ‘*‘Color 
brown, densely freckled with blackish brown. 
Soft dorsal with a pale blotch in the middle of 


its anterior half, caudal with a broad pale band 
across the centre flecked with dark brown. Pec- 
toral with a pale central band becoming more 
distinct towards its upper edge. A white spot in 
front of each orbit.’’ Preserved specimens about 
as in Figure 4. Darkest below front, middle, 
and end of spinous dorsal fin; color extend- 
ing onto fins. Soft dorsal fin darkest distally. 
Caudal fin streaked with brown, darkest just be- 
yond middle, also a dark patch near base. Inside 
of pectoral fin as in Figure 1g, h; two dark areas, 
with streaks of darker brown in some speci- 
mens; inside nearly uniform dark brown in one 
[discolored?] specimen (ANSP 98707). Color- 
ation on inner surface of pectoral fin shows 
through from outside of fin. 

Measurements for seven specimens (86.8—170 
mm SL) in percent SL: head 34-36, snout 15— 
17, orbit 6-7, interorbital width 7-9, postorbital 
12-14, pectoral fin 38-48. 

DISTRIBUTION.—Inimicus caledonicus 1s 
known from New Caledonia, Queensland coast 
of Australia, and the Andaman and Nicobar is- 
lands in about 15 to 60 m. A wider distribution 
is expected. 


Inimicus didactylus (Pallas) 
(Figures li-l, 5, 8-10; Table 1) 


Scorpaena didactyla PALLAS, 1769:26-29, pl. 4, figs. 1-3 
(original description; no type-locality). SHAW 1803:279-280 
(compiled). 

Trigla rubicunda HORNSTEDT, 1788:49-51, pl. 3 (original de- 
scription; type-locality Amboina; figure poor). 

Synanceia didactyla: BLOCH AND SCHNEIDER 1801:195-196 
(compiled description and distribution). 

Synanceia rubicunda; BLOCH AND SCHNEIDER 1801:196 (com- 
piled). 

Pelor maculatum CUVIER in CUVIER AND VALENCIENNES, 
1829:434-435 (original description; type-locality Waigiou). 
Lesson 1830:210, pl. 20 (Waigiu). CUvIER 1834:27, fig. 2 
(figure poor). BLEEKER 1849:3—-4 (listed; Waigioe); 1851:20, 
22 (listed; Waigioe); 1852b:242 (listed; Waigioe). GUNTHER 
1860:150 (compiled from Cuvier 1829). BLEEKER 1873:141 
(listed; Amoy, China). PETERS 1876b:834 (listed; New Brit- 
ain). BLANC AND HUREAU 1968:27 (type at MNHN). 

Pelor obscurum CUVIER in CUVIER AND VALENCIENNES, 
1829: 436-437 (original description; type-locality Port Pras- 
lin, New Ireland). LEsson 1830:211, pl. 21, fig. 2 (New Ire- 
land). BLEEKER 1854b:360 (listed; Batjan). SAUVAGE 
1878: 148 (compared with P. caledonicum). BLANC AND Hu- 
REAU 1968:28 (type at MNHN). 

Scorpaena digitata GRAY, 1854:117—118 (original description; 
type-locality ‘‘Indian seas’’). 

Pelor didactylum: GUNTHER 1860:150 (compiled; brief de- 
scription; synonymy). SAUVAGE 1878:147 (compared with 
P. caledonicum). SEALE 1906:83 (listed; one specimen from 
Fate, New Hebrides; BPBM 963). PELLEGRIN 1912:206 
(Port Sandwich, New Hebrides). WEBER 1913:497-498 (syn- 


490 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


FIGURE 5. 


onymy; one specimen from SIBOGA sta. 258, Kei Is., in 22 
m). FOWLER 1928:298-299 (synonymy; description; one 
specimen from Fate, New Hebrides). HERRE 1931: 13 (list- 
ed; New Hebrides). FOWLER 1938c:201 (brief synonymy; 
Singapore). 

Pelor didactylus: BLEEKER 1861:49 (listed; Singapore); 
1873:141 (listed; China); 1874:4, 7-8, pl. 4, figs. l-la (de- 
scription; good figures; synonymy; [p. 22, list of figures in 
error)). 

Simopias didactylus: GiLv 1905:221 (Synanceja didactyla and 
S. rubicunda as treated by Bloch and Schneider as syn- 
onyms of Simopias didactylus). 

Inimicus didactylum: FOWLER 1918:65 (listed; new record for 
Philippines). 

Inimicus didactylus: FOWLER 1927:288—289 (description; one 
specimen from Philippines). HERRE 1934:66 (nine specimens 
from the Philippines; smallest one with 17 dorsal spines). 
MATSUBARA 1943:418—420, fig. 143 (synonymy; detailed de- 
scription based on a specimen from Pelew I.). DE BEAU- 
FORT in WEBER AND DE BEAUFORT 1962:104—106, fig. 30 
(synonymy; good description; distribution). McKay 1964:10 
(compared with Inimicus barbatus and I. cirrhosus). TorR- 
TONESE 1964:79 (listed; two specimens from Korido, New 
Guinea). 

Inimicus bifilis FOWLER, 1938a:85—86, fig. 38 (original descrip- 
tion; type-locality Canmahala Bay, Philippines, ALBA- 
TROSS). McKay 1964:11 (suggested J. bifilis might be a ju- 
venile of J. didactylus or I. cuvieri). 


REMARKS.—The nominal species Trigla rubi- 
cunda Hornstedt, Pelor maculatum Cuvier, Pe- 
lor obscurum Cuvier, and Scorpaena digitata 
Gray all have been previously referred to didac- 
tylus. McKay (1964) suggested that /. bifilis was 
a juvenile of J. didactylus or I. cuvieri; our ex- 


Inimicus didactylus, WAM P5801, 113 mm SL, New Hebrides. 


amination of the specimen shows that it is a ju- 
venile of J. didactylus. 


MATERIAL EXAMINED.—RMNH 669 (104, holotype of 
Scorpaena didactyla), no specific locality given. 

Thailand: CAS 15165 (1, 136). 

Singapore: CAS 15159 (1, 110). NMS 208 (1, 107). 

South Viet Nam: CAS 15166 (1, 94.3). 

Indonesia: ZMA 101.829 (1, 102), Java Sea. ZMA 101.900 
(1, 84.9), Kei Is. ZMA 109.799 (1, 43.7), Kei Is. 

Philippines: USNM 98905 (40.7, holotype of Inimicus bifi- 
lis), Canmahala Bay, ALBATROSS, 11 Mar. 1909. CAS 15164 
(1, 84.6). NMP 2042 (1, 90.5). NMP 5218 (1, 101). SU 28360 
(4, 46.6-116). SU 28361 (2, 111-120). SU 29785 (1, 88.1). 
USNM 168173 (1, 29.4). USNM 136457 (1, 80.6). USNM 
136458 (1, 79.8). USNM 136459 (1, 33.9). USNM 136460 (1, 
44.8). 

Northern Australia: CAS 31426 (1, 41.5), Timor Sea. 

New Guinea: MNHN 6726 (130, holotype of Pelor macula- 
tum), Waigiou, Lesson and Garnot. MNHN 718 (1, 87.3). 
MNHN 6734 (1, 112). MNHN 6735 (1, 81.0). USNM 218504 
(1, 132). USNM 218535 (1, 61.0). 

Palau Is.: CAS 15160 (3, 120-149). CAS 15161 (1, 116). CAS 
15162 (1, 117). CAS 15163 (2, 97.7-136). 

Bismark Archipelago: MNHN 6725 (125, holotype of Pelor 
obscurum), New Ireland, Lesson and Garnot. MCZ 25150 (1, 
144), New Britain. 

Solomon Islands: USNM 144298 (1, 88.2). 

New Hebrides: BPBM 963 (1, 103). MNHN 94-364 (1, 120). 
WAM P5801 (1, 113). 


DIAGNosIs.—No filamentous pectoral rays in 
adults. Orbits moderately far apart, not highly 
elevated. Snout longer than postorbital distance 
(postorbital distance into snout 1.2-1.7). In- 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 49] 


FIGURE 6. 


Inimicus brachyrhynchus 


terspinous membranes from fourth dorsal spine 
rearwards less than one-third of spine height. 
Inner pectoral fin coloration (Figs. li-l) diag- 
nostic. 

DESCRIPTION.—Dorsal fin with 15S—17 (usually 
16) spines and 7-9 (usually 8 or 9) soft rays, total 
23-26 (usually 24 or 25). Anal fin with 2 spines 
and 10—12 (usually 11) soft rays, total usually 13. 
Upper | or 2 pectoral rays filamentous in juve- 
niles under about 50 mm SL (to 46.3 mm SL in 
our material). Gill rakers on outside of first arch 
total 9-11, 1-3 on upper arch, 7—9 on lower arch. 
Vertebrae 28 (5 specimens). Head spines and 
general features as for genus. 

General body shape and coloration of pre- 
served specimens as in Figure 5. In preserva- 
tive, body and head brown mottled with paler 
areas, variable. Underparts tan with darker ir- 
regular spots or tan with pale spots. White spots 
in area before eyes and in occiput not well 
marked. Interspinous membrane of spinous dor- 
sal dark brown, especially between spines 1-3. 
Soft dorsal fin pale in middle. Caudal fin with 
dark band near base and a broad subterminal 
band. Anal fin mostly dark. Pelvic fins dark 
brown to blackish, usually streaked with white 
near base. Pectoral fin usually with a pale area 
at base, followed by large dark area, a pale band, 
then dark distally. Inner surface of pectoral fin 
distinctive (Figs. li-l); streaks of white some- 
times as roundish pale patches. 


, CAS 31424, 109 mm SL, Hong Kong. 


Measurements for 32 specimens (43.7—148 
mm SL) in percent SL: head 32-37, snout 15— 
20, orbit S—6, interorbital width 5—11, postorbital 
10-15, pectoral fin 39-48. 

DISTRIBUTION.—Inimicus didactylus occurs 
from the Indo-Australian Archipelago to the 
Philippines, Micronesia, and Melanesia. 


Inimicus brachyrhynchus (Bleeker) 

(Figures Im, 6, 8; Table 1) 

Pelor brachyrhynchus BLEEKER, 1874a:4-7, pl. 2, fig. 4 and 
pl. 3, fig. 2 (original description; type-locality Singapore; 
fairly good figure). BLEEKER 1878:fig. 1 on pl. 413 (figure 
from Bleeker 1874; [fig. Id of pectoral fin labeled as P. 
brachyrhynchus is P. cuvieri)). 

Pelor cuvieri: BLEEKER 1878:pl. 416, fig. 3a ({[switched fig. la 
on pl. 413 with fig. 3a on pl. 416 J; from Bleeker 1874). 
Inimicus brachyrhynchus: DE BEAUFORT in WEBER AND DE 
BEAUFORT 1962:105—107 (description of holotype and one 
additional specimen with no data). McKay 1964:10 (com- 

parison with /. cirrhosus). 


MATERIAL EXAMINED.—RMNH 5907 (130, holotype of /. 
brachyrhynchus), Singapore, P. Bleeker. CAS 31424 (1, 109), 
Hong Kong. 

DIAGNOosIS (based on two specimens).—No 
filamentous pectoral rays in adults. Orbits only 
slightly elevated. Snout shorter than postorbital 
distance (postorbital distance into snout 0.7— 
(0.8). Interspinous membranes from fourth dorsal 
spine rearwards extend up to about one-half of 
spine height. Inner pectoral fin coloration diag- 
nostic (Fig. Im). 

DESCRIPTION.—Dorsal fin with 17 spines and 


492 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


8 branched soft rays, total 25. Anal fin with 2 
spines and 10 soft rays, total 12. Gill rakers on 
outside of first arch total 11-13, 3 on upper arch, 
8-10 on lower arch. Vertebrae 29 (1 specimen). 
Head spines and general features as for the ge- 
nus. 

General body shape and coloration of pre- 
served specimens as in Figure 6. Head and body 
mostly dark brownish, with belly paler. Inter- 
spinous membrane with white patches between 
about spines 3—5 and 13-17; pale areas extend- 
ing on to body. Slightly oblique pale patch along 
anterior part of soft dorsal fin extending about 
half way down body. Caudal fin with pale areas. 
Inner surface of pectoral fin as in Figure Im. A 
transverse white band at about middle of pec- 
toral fin as seen from outer surface. On inner 
surface band made up of large white spots. 

Measurements for 2 specimens (109 and 130 
mm SL) in percent SL: head 33-35, snout 12- 
13, orbit 5, interorbital width 9-10, postorbital 
16-17, pectoral fin 32-35. 

DISTRIBUTION.—lnimicus brachyrhynchus is 
known only from two localities, Singapore and 
Hong Kong. A wider distribution is expected. 


Inimicus joubini (Chevey) 

(Figure In, 7a, 8; Table 1) 

Chorismopelor joubini CHEVEY, 1927:222-224, fig. (original 
description; type-locality Tonkin Gulf, Haiphong, figure 
poor; comparisons with other genera; as type of genus 
Chorismopelor). 

Chorismopelor joubini: BLANC AND HUREAU 1968:26 (type at 
MNHN). 


REMARKS.—Chevey distinguished this species 
from those of the genus Jnimicus principally on 
the basis of its having three instead of two free 
lower pectoral rays. The third ray from the bot- 
tom is free distally, more so than in the other 
species of Jnimicus, but the third ray is much 
more closely joined to the rays above than to 
the lower two free rays (Fig. In). In all other 
features joubini seems to be closely related to 
species of the genus Jnimicus, particularly the 
Japonicus subgroup, and we do not feel this 
small difference is of either generic or subge- 
neric importance. We have located only one ad- 
ditional specimen that we feel is J. joubini, al- 
though its third pectoral ray from the bottom is 
free for only about one-sixth of its length. 


MATERIAL EXAMINED.—MNHN 26.398 (158, holotype of 
C. joubini), Haiphong, Tonkin Gulf, North Vietnam, P. Chev- 
ey. SU 67600 (1, 111), Hiroshima, Japan, D. S. Jordan and J. 
O. Snyder. 


DIAGNOsIS (based on two specimens).—No 
filamentous pectoral rays in adults. Third pec- 
toral ray from the bottom free for about one- 
third its length as in Figure In. Orbits slightly 
elevated. Snout shorter than or nearly equal to 
postorbital distance (postorbital distance into 
snout 0.7-0.9). Interspinous membranes from 
fourth dorsal spine rearwards extend up to one- 
half spine height. Inner pectoral fin coloration 
(Fig. In) mostly plain colored, not strikingly 
marked. 

DESCRIPTION.—Dorsal fin with 17 spines and 
7 soft rays, total 24. Anal fin with 2 spines and 
8-9 soft rays, total 10-11. Lower two pectoral 
rays free; third ray from bottom free for part of 
its length (about one-third its length in type and 
one-sixth in second specimen). Gill rakers on 
outside of first arch total 9, 2-3 on upper arch, 
6-7 on lower arch. Vertebrae 28 (2 specimens). 
Head spines and general features as for genus. 

General body shape and coloration of pre- 
served specimens as in Figure 7a. Color of ho- 
lotype blackish brown with dark spots and 
markings over body and head. White patches 
between dorsal spines 3-4 and 8-11, pale areas 
extending on to body. Fins with variegated pat- 
tern. Second specimen light colored (Fig. 7a). 
Inner pectoral coloration as in Figure In; back- 
ground pale with light and dark patches similar 
to that of J. japonicus. 

Measurements for 2 specimens (110-158 mm 
SL) in percent SL: head 35-36, snout 13-15, 
orbit 5—6, interorbital width 8-10, postorbital 
15-18, pectoral fin 37-42. 

COMPARISONS.—Inimicus joubini is very sim- 
ilar to J. japonicus in most features, including 
counts and shape. The coloration on the inner 
surface of the pectoral fin also is similar. [nim- 
icus joubini appears to have the area at the front 
of the dorsal fin slightly more elevated than in 
I. japonicus. In nearly all of our specimens of 
I. japonicus, the third ray from the bottom is 
fully joined to the ray above, with the membrane 
extending all the way to the tip; in a few speci- 
mens the tip is barely free. In /. joubini the third 
ray from the bottom is free for part of its length 
as discussed above. Inimicus brachyrhynchus, 
the third species in this species group as dis- 
cussed under the genus, can be separated on the 
basis of pectoral fin coloration as given in the 
key. 

DISTRIBUTION.—Inimicus joubini is known 
from the holotype from the Tonkin Gulf, North 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 493 


FIGURE 7. 


Viet Nam, and from another specimen from Ja- 
pan. 


Inimicus japonicus (Cuvier) 
(Figures lo, 7b, 8; Table 1) 


Pelor japonicum CUuviER in CUVIER AND VALENCIENNES, 
1829:437, pl. 93 (original description; type-locality Japan). 
TEMMINCK AND SCHLEGEL 1843:44, pl. 18, fig. 2 (descrip- 
tion; Nagasaki, Japan). RICHARDSON 1846:212 (short de- 
scription; distribution, Japan and China). GUNTHER 
1860:151 (Pelor tigrinum in synonymy; short description; 
China and Japan). STEINDACHNER AND DODERLEIN 
1884:197 (Pelor auranticum in synonymy; compared with 
P. japonicum; Tokyo). NySTROM 1887:19 (listed; Nagasaki, 
Japan). STEINDACHNER 1896:203 (short description; Kobe, 
Japan). ISHIKAWA AND MATSUURA 1897:49 (listed, Japan). 
JORDAN AND SNYDER 1901a:367 (listed; Tokyo). JORDAN 
AND SNYDER 1901b:100 (listed; Yokohama, Nagasaki, and 
Shimoda, Japan). 


(a) Inimicus joubini, SU 67600, 111 mm SL, Hiroshima, Japan; (b) J. japonicus, UT 51839, 190 mm, Japan. 


Pelor auranticum SCHLEGEL in TEMMINCK AND SCHLEGEL, 
1843:44, pl. 18, fig. 1 (original description; Nagasaki, Ja- 
pan). RICHARDSON 1846:212 (listed; Japan). GUNTHER 
1860: 151 (compiled from Schlegel 1843). JoRDAN AND SNyY- 
DER 1901b:100 (listed; Nagasaki and Kaminoseki, Japan). 

Pelor tigrinum RICHARDSON, 1846:212 (original description; 
type-locality Canton, China). 

Inimicus auranticus: JORDAN AND STARKS 1904:160 (key; de- 
scription; Hiroshima and Kobe, Japan). 

Inimicus japonicus: JORDAN AND STARKS 1904:159 (key; de- 
scription, southern Japan). FRANZ 1910:74 (listed; Sagami- 
bai and Aburatsubo, Japan). JORDAN AND METz 1913:52 
(listed; Fusan, Korea). JoRDAN, TANAKA, AND SNYDER 
1913:249 (listed; Pelor auranticum included as a possible 
synonym; Tokyo to Nagasaki, Japan). JORDAN AND 
THOMPSON 1914:276 (listed; Misaki, Japan). JORDAN AND 
Husss 1925:275 (listed many Japanese localities; common 
in the Japanese markets southwards). Mort 1928:7 (listed; 
Oni-okoze and Ruganho, Japan). SCHMIDT AND LINDBERG 
1930: 1146 (listed; Tsuruga Bay, Japan). SCHMIDT 1931a:111 


494 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


_20” 40° 60° 8go° 100° 120° 140° I60° I80° 


mul tibarbus 
natalensis 
. caledonicus 


. filamentosus 


a — el — a o> Te om) 


. Sinensis 
20° 2 


o° 


20° 


40° 


20° 40° 60° 8go° 100° (20° 140° 160° Ig0° 
o 


20° 


o° 


20° . brachyrhynchus 
. cuvieri 


. didactylus 


. japonicus 


. joubini 


40° 


FiGureE 8. Distribution of the species of the genera Choridactylus and Inimicus. 


(listed; Nagasaki and Misaki, Japan and Fusan, Korea); onymy; good description; internal features; Japanese spec- 
1931b:116 (listed; Nagasaki and Obama, Japan). TANAKA imens; distribution). Mort 1952:156 (listed; Pusan, Quelpart 
1931:36 (listed; Japan). KAMOHARA 1938:49 (listed; Okoze, I., South Korea, and Yonganpo, Japan). Moon Ki 1956:27 
Japan). TORTONESE 1939:361—362 (description; color vari- (listed; southern Japan). Mort 1956:27 (listed; southern Ja- 


ation; Yokohama, Japan). MATSUBARA 1943:415—-418 (syn- pan). 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 495 


Inimicus japonicus var. auranticus FRANZ, 1910:74 (descrip- 
tion of three color varieties in Japan). 

Scorpaenopsis cirrhosa: HALSTEAD 1970:pl. XXVIII (3 color 
phases). 


MATERIAL EXAMINED.—Hong Kong: CAS 27953 (1, 162) 
FRSHK uncat. (1, 145). MCZ 13977 (2, 184-186). SU 29256 
(1, 212). SU 60958 (1, 180). 

China: CAS 868 (1, 100). MCZ 13975 (1, 103). USNM 
130535 (1, 174). USNM 130596 (2, 155-178). 

Korea: CAS 15769 (2, 139-191). CAS 31926 (1, 126). MCZ 
47707 (2, 48.5—189). SU 25279 (1, 140). 

Japan: RMNH 597 (235, stuffed specimen, lectotype of Pe- 
lor auranticum), Burger collection. RMNH 614 (166, paralec- 
totype of auranticum), Von Siebold collection. CAS 11245 (1, 
143). CAS 31435 (1, 136). CAS 31436 (1, 158). CAS 31434 (1, 
155). CAS 31433 (1, 129). SU 3698 (2, 79.1-168). SU 7170 (1, 
217). SU 7179 (1, 176). SU 7417 (1, 204). SU 7851 (2, 81.1- 
162). SU 23560 (2, 200-206). UT 51839 (1, 190). Additional 
material is present in the USNM. 


DIAGNosIs.—No filamentous pectoral rays in 
adults. Orbits somewhat elevated. Snout shorter 
than postorbital distance (postorbital distance 
into snout 0.7—-0.9). Interspinous membrane 
from fourth dorsal spine rearwards extends up 
to one-half of spine height. Inner pectoral fin 
coloration variable (usually as in Fig. lo), with 
no diagnostic pattern. 

DESCRIPTION.—Dorsal fin with 16—18 (usually 
17) spines and 5-8 (usually 7) soft rays, total 23— 
25 (usually 24). Anal fin with 2 spines and 9-10, 
rarely 8 soft rays, total 10-12. Upper pectoral 
ray filamentous in 48.5-mm-SL specimen; not 
filamentous in larger specimens. Gill rakers on 
outside of first arch total 10-12, 2-4 on upper 
arch, 7-9 on lower arch. Vertebrae 28 (2 speci- 
mens) and 29 (1). Head spines and general body 
features as for the genus. Body with many skin 
appendages, particularly on head, and on dorsal 
and pectoral fins. 

General body shape and coloration of pre- 
served specimens as in Figure 7b. In general, 
color of body highly variable. Head and body 
mostly brown, underparts paler; body often light 
colored except for dark spots on underparts. 
Head with variable dark patches. Broad pale 
patches often present between dorsal spines 3- 
5, 11-14, and beginning of soft dorsal, pale areas 
extending along the lateral side of body. Fins 
variously blotched with paler color on dark 
background. Pectoral and caudal fins with white 
and brown bands. Inner surface of pectoral fin 
as in Figure lo, variable, without a conspicuous 
pattern, most often streaked or spotted brown; 
in some dark gray to light brownish, with black 
spots or patches. 


Color in life variegated, often reddish or 
brown mottled with white. Sometimes mostly 
yellow with dark spots (“‘auranticus’’ pattern) 
[see Halstead 1970:pl. LX for three color vari- 
ations]. 

Measurements for 28 specimens (75.3-217 
mm SL) in percent SL: head 29-44, snout 12- 
15, orbit 4-6, interorbital width 8-11, postorbital 
16-19, pectoral fin 3546. 

DISTRIBUTION.—Inimicus japonicus is an 
Oriental species occurring from southern Japan 
to Hong Kong. 


DISTRIBUTION AND REMARKS ON BIOLOGY 


Members of the subfamily Choridactylinae 
occur in the Indo-West Pacific (Fig. 8), from Ja- 
pan southward throughout the Indo-Australian 
Archipelago to New Caledonia and Australia, 
and westward to the western Indian Ocean and 
Red Sea. Most species have fairly limited 
ranges. Members of this subfamily are not found 
in the central Pacific and represent another ex- 
ample of a group that seemingly could, but does 
not cross the Andesite Line marking the sepa- 
ration of continental and oceanic rocks (see 
Springer and Goman 1975:16). They occur on 
open sand or silty substrate (McKay 1964). 
Depths of capture range from near shore (about 
5 m) to about 90 m, with most species found 
shallower than about 40 m. 

Little is known about the biology of these fish- 
es. They appear to be opportunistic predators 
feeding mostly on other fishes. They have ven- 
omous fin spines and erect the dorsal spines 
when aroused, as do many other scorpionfishes. 
They also have bright colors on the inner surface 
of the pectoral fin (see Condé 1977), and the fins 
are spread outward and rolled forward when the 
fish is aroused, thereby creating a flash of bright 
color from an otherwise drab individual; pre- 
sumably this startles would-be predators. This 
behavioral response appears in some other scor- 
pionfishes, occurring in such subfamilies as the 
Scorpaeninae and probably the Minoinae, and 
perhaps other subgroups. It is unknown whether 
this behavioral response is used in interspecific 
interactions. 

Morphologically, a number of interrelated 
modifications and trends in the subfamily are 
directed towards settling into the substrate, con- 
cealment, and feeding behavior: 

1. The lowermost two (Unimicus) or three 


496 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 21 


FIGURE 9. 


(Choridactylus) pectoral fin rays are free of 
membrane (see Fig. 9). The free rays are used 
in ‘‘walking’’ on the bottom. The osteology of 
the pectoral girdle has been described by Samuel 
(1961). 

2. The gas bladder is lost, but a drumming 
muscle remains (see Hallacher 1974:73). 

In the genus /nimicus, the following additional 
modifications are found: 

1. The eyes become situated high up on the 
head, well above the dorsal profile in the more 
specialized species (see Fig. 9). This would ap- 
pear to permit deeper burying in the substrate. 

2. The membranes of the dorsal fin, from the 
fourth spine rearward, become deeply incised. 
This is most advanced in species such as /. fil- 
amentosus (see Condé 1977; fig. 4) and J. didac- 
tylus (figs. 9-10). This reduction in membrane 
allows the spines to be folded laterally (see fig. 
10). Skin flaps on the spines contribute to con- 
cealment. 

3. The anterior three dorsal spines form an 


Live adult Inimicus didactylus in prowling posture. Note position of two free lower pectoral fin rays, location 
of eye, and anterior dorsal spines. (Photo by Tom McHugh in Steinhart Aquarium.) 


apparent lure, reminiscent of that found in /ra- 
cundus signifer (see Shallenberger and Madden 
1973), but our aquarium observations of J. di- 
dactylus indicate that the lure is stationary and 
not moved as in Jracundus. 

4. The mouth is more upturned in more ad- 
vanced species of /nimicus, presumably allow- 
ing deeper settlement in the substrate. 

As observed for J. didactylus in Steinhart 
Aquarium, settling into the sand is accomplished 
by a rocking of the body and pushing of sand 
from underneath the body by the large pelvic 
fins. The lowermost two free pectoral rays are 
extended forward (Fig. 9), and they are used to 
pull the body out of the substrate when striking 
prey. Figure 9 shows the posture just before 
feeding. Engulfment of prey is extremely rapid. 
Inimicus didactylus was observed to walk on the 
bottom of its free pectoral rays while ‘‘stalking”’ 
prey and moving around the tank. Additional 
observations are provided by Condé (1977) for 
I. filamentosus. 


ESCHMEYER, RAMA-RAO, & HALLACHER: CHORIDACTYLINE SCORPIONFISHES 497 


FiGuRE 10. 


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PROCEEDINGS 


OF THE 


CALIFORNIA ACADEMY OF SCIENCES 


FOURTH SERIES 


Vol. XLI, No. 22, pp. 501-528; 14 figs. 


March 2, 1979 


SYNOPSIS OF THE TURTLES, CROCODILES, AND 
AMPHISBAENIANS OF IRAN 


By 


Steven C. Anderson! 


Raymond/Callison College and Department of Biology, 
University of the Pacific, Stockton, California 95211 


ABSTRACT: 


Three species of freshwater turtles, Emys orbicularis, Mauremys caspica caspica, Trionyx 


euphraticus; two land tortoises, Agrionemys horsfieldii, Testudo graeca (T. g. ibera and T. g. zarudnyi), 
and one marine turtle, Eretmochelys imbricata bissa, are known and documented from Iran. Chelonia 
mydas japonica and Dermochelys coriacea have been observed, and two other species of marine tugtles, 
Caretta caretta gigas and Lepidochelys olivacea, may occur in Iranian waters. A single species of crocodile, 
Crocodylus palustris, and a single amphisbaenian, Diplometopon zarudnyi, occur in Iran. Synonymy, di- 
agnoses, Iranian distribution, all literature records, and all known Iranian localities are given, and the natural 
history and ecology of each species in Iran is summarized. A key to the turtles of Iran is provided. 


INTRODUCTION 


This synopsis is an outgrowth of work begun 
in 1958 with field studies in southwestern Iran, 
continued during subsequent years through 
studies of museum collections, and culminated 
with field studies throughout Iran in 1975. 

With the encouragement and sponsorship of 
the Iran Department of the Environment, a num- 
ber of ecosystem studies have been initiated in 
various areas of Iran, and environmental impact 
statements are now being undertaken in connec- 
tion with development projects. It is thus desir- 
able that synopses of the various taxonomic 
groups of organisms be prepared whenever pos- 
sible to facilitate these studies. The systematics 
of amphibians and reptiles has now reached the 
stage where such comprehensive summaries are 


1 Research Associate, Department of Herpetology, Califor- 
nia Academy of Sciences; Research Associate, Iran Depart- 
ment of the Environment. 


possible. The present report is but a small part 
of continuing systematic work on the herpetol- 
ogy of Iran, but a single monographic treatment 
of all of the amphibians and reptiles has not 
proved practical nor economically feasible. 
Therefore, reports on smaller taxonomic groups 
are being published as this work reaches the ap- 
propriate stage for each group. 

The turtles of Iran are representative of 
the impoverished chelonian fauna of the Pa- 
learctic. Emys, Testudo, and the western forms 
of Mauremys today are all restricted in their 
distribution to the milder climates of southern 
Europe, North Africa, and Southwest Asia, 
being essentially Mediterranean faunal ele- 
ments. Agrionemys may be a relatively ancient 
genus of uncertain affinities; its distribution is 
strictly Irano-Turanian. The marsh crocodile, 
which occurs in the extreme southeastern corner 
of Iran, is one of a small number of Indian faunal 
elements extending beyond the Oriental Region 


{501] 


502 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


into Iran. In the not-too-distant past, the cur- 
rently interrupted distribution of crocodiles in 
the river systems of the Makran of Iran and Paki- 
stan may have been more or less continuous, 
connecting the Iranian population with those of 
the Hab and Indus drainages. In southwestern 
Iran, in the low plains of Khuzestan, Arabian 
elements of the Saharo-Sindian subregion are 
encountered, and Iran’s lone amphisbaenian, 
Diplometopon zarudnyi, is one of these. It is the 
northeasternmost representative of the Trogo- 
nophidae, amphisbaenians occurring in the arid 
regions of North Africa, East Africa, and the 
Arabian peninsula. 

In the accounts which follow, standard her- 
petological procedures and terminology have 
been followed. The reader not familiar with her- 
petological terminology is referred to Peters 
(1964) for definitions. 

The synonymies include all references known 
to me that cite specimens from Iran. Thus, my 
interpretations of the identifications of previous 
workers and the status of all published museum 
records should be clear. 

I have attempted to summarize what little is 
known of the natural history of Iranian species, 
based on published accounts and my own ob- 
servations. Where no information exists for a 
species within Iran, I have cited accounts for 
areas elsewhere in the range of the species. 

I have listed here all of the Iranian localities 
known for each species. Wherever possible, I 
have associated the localities in the literature 
with a standardized spelling in the United States 
Board on Geographic Names Gazetteer no. 19, 
Iran, 1956, and only this spelling has been listed 
under each species in the interest of saving 
space. Each published locality is followed by the 
initials of the author and the date, and/or (in the 
case of specimens I have examined) the abbre- 
viation for the museum in which the specimen(s) 
is located. Where I have been unable to asso- 
ciate a published place name with a standardized 
spelling, the original spelling has been used. Any 
localities from which I have examined speci- 
mens are preceded by an asterisk (*). In cases 
where the reader questions my interpretation of 
the identification by a previous author, it some- 
times may be necessary to check the locality 
cited in the original paper. Distances and ele- 
vations appear as given in publications and on 
museum labels. In fear of introducing further im- 
precision, I have not converted such data from 


miles and feet to kilometers and meters. For 
convenience in locating place names, the local- 
ities have been listed under the ostdn (admin- 
istrative province) in which they occur. A map 
of these ostan boundaries is provided in a pre- 
vious paper (Anderson 1974: 28, fig. 1), and in 
Fisher (1968: map 2). It should be noted, how- 
ever, that the boundaries of administrative sub- 
units in Iran have changed since publication of 
this map, and seem to be subject to periodic 
relocation. It seems practical to stabilize this 
organization of localities for expedient refer- 
ence, however. 


ACKNOWLEDGMENTS 


Anyone engaged in faunal studies owes his 
primary debt to the many collectors, without 
whom such studies would be impossible. Many 
of these collectors are anonymous, but the bulk 
of Iranian material has been collected by those 
cited in the publications of previous workers. I 
Owe special thanks to William S. and Janice K. 
Street, who have contributed so much and have 
waited so long and patiently for the reports on 
their material. Similarly, Dr. Henry Field has 
been a tireless contributor to the natural history 
material of Southwest Asia, and has constantly 
encouraged those of us working with the fauna. 

Curators who have loaned me material and/or 
made me welcome at their institutions during the 
course of my work include: Robert Inger and 
Hymen Marx, Field Museum of Natural Histo- 
ry; the late Doris Cochran, the late James A. 
Peters, and George Zug, United States National 
Museum; Charles M. Bogert, Richard Zweifel, 
and Charles Myers, American Museum of Nat- 
ural History; Charles Walker, Donald Tinkle, 
and Arnold Kluge, Museum of Zoology, Uni- 
versity of Michigan; Ernest E. Williams, Mu- 
seum of Comparative Zoology, Harvard Uni- 
versity; Robert C. Stebbins, Museum of 
Vertebrate Zoology, University of California, 
Berkeley; Wilmer W. Tanner, Brigham Young 
University; Ilya Darevsky, Zoologicheski’i In- 
stitut, Leningrad; Josef Eiselt, Naturhistorisch- 
es Museum, Wien; Alice Grandison, J. C. Bat- 
tersby, and E. N. Arnold, British Museum 
(Natural History); Jean Guibé, Muséum Nation- 
al d’ Histoire Naturelle, Paris; F. W. Braestrup, 
Universitetets Zoologiske Museum, Copenha- 
gen; P. Kuenzer, Zoologische Institut und Mu- 
seum der Universitat, GOttingen; Robert G. 


ANDERSON: IRANIAN REPTILES 


Tuck, Jr., Iran National Museum of Natural 
History (Muse’-ye Melli-ye Tarikh-e Tabi’1), 
Tehran. 

Much of the research for this and other studies 
in Southwest Asian herpetology was done while 
I was Associate Curator in the Department of 
Herpetology of the California Academy of Sci- 
ences. I thank Dr. George E. Lindsay, Director 
of that institution, and Dr. Alan E. Leviton, 
Chairman and Curator of the Department of 
Herpetology for the opportunity and necessary 
facilities to pursue the research. Dr. Leviton and 
I have worked together on many projects in 
Southwest Asian herpetology, and his com- 
ments and encouragement have been a constant 
stimulus. 

During field work in 1958, I was extended 
many courtesies by employees of the Iranian Oil 
Exploration and Producing Company. The Iran 
Department of the Environment made possible 
my field work in 1975 as a visiting scientist, and 
provided me with air travel funds, vehicles, 
equipment, and field assistants. I am indebted 
to many personnel of that department, but es- 
pecially to His Excellency, Mr. Eskandar Fi- 
rouz, the former Director, and to Robert G. 
Tuck, Jr., Curator and Advisor to the Museum. 

I have been accompanied in the field by many 
people at various times. I am particularly grate- 
ful to Howard T. Anderson, William O. Wil- 
liams, and Rufus Cook, all formerly with the 
Iranian Oil Exploration and Producing Compa- 
ny; Ruben McCullers, American Peace Corps; 
Reza Khazai’e, Abbas Ahmadi, and Robert G. 
Tuck, Jr., Iran Department of the Environment. 

Portions of the work were supported by a 
grant from the American Philosophical Society 
(Grant no. 4959—Penrose Fund, 1968), by three 
Faculty Research Grants (1971, 1972, 1975) from 
the University of the Pacific, and by travel and 
field assistance from the Iran Department of the 
Environment (1975). The University of the Pa- 
cific made possible the 1975 field work through 
a faculty development leave. 

Most important to me has been the continuing 
support of my family. My father, Howard T. 
Anderson, has provided financial assistance as 
well as constant encouragement and intellectual 
stimulation throughout the 20 years of my her- 
petological work. My mother, Lois B. Ander- 
son, shared her home in Masjed Soleyman with 
elements of the Iranian herpetofauna during 
1958. My wife, Kay, and son, Malcolm, have 


503 


) | 


Sa-G 


Ficure 1. (A) Dermochelys. (B) Trionyx. 


made all of the sacrifices familar to families of 
researchers. 


SYSTEMATIC SECTION 


The following abbreviations are used for mu- 
seums: AMNH—American Museum of Natural 
History, New York. CAS—California Academy 
of Sciences, San Francisco. FANH—Field Mu- 
seum of Natural History, Chicago. MMTT— 
Muze’-ye Melli-ye Tarikh-e Tabi’i, Tehran. 
USNM—United States National Museum, 
Washington, D.C. 


504 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


FIGURE 2. 
m = marginal, n = nuchal, v = vertebral. 


Key to the Turtles of Iran 


Species marked with an asterisk (*) have not 
been collected in Iran, but may be expected to 
occur within the borders of the country. De- 
partment of the Environment personnel have re- 
corded turtles identified as Chelonia mydas and 
Eretmochelys imbricata from many areas along 
the Iranian coast of the Persian Gulf and the 
islands of the gulf. Ten to 15 Dermochelys cor- 
iacea are reportedly taken annually at Larak Is- 


(A) Carapace of Eretmochelys. (B) Carapace of Chelonia. (C) Carapace of Caretta. Abbreviations: c = costal, 


land in the Strait of Hormoz to support a local 
oil industry for sealing the hulls of dhows and 
other craft. But there are no records of marine 
turtles in the herpetological literature, and apart 
from two skulls of Eretmochelys imbricata bis- 
sa, no specimens of record. 


la. Carapace and plastron without horny 


plates: (Bis. 1)... 2 
1b. Carapace and plastron with horny plates 
(Pigs .2=4) << 3. 2 ee 3 


FIGURE 3. 
g = gular, h = humeral, i = inframarginal, p = pectoral. 


(A) Plastron of Caretta. (B) Plastron of Lepidochelys. Abbreviations: ab = abdominal, an = anal, f = femoral, 


ANDERSON: IRANIAN REPTILES 


2a. 


2b. 


3a. 


3b. 


4a. 


4b. 


Dae 


Sb. 


6a. 


6b. 


TA. 


7b. 


8a. 


8b. 


9a. 


9b. 


Carapace with 7 distinct longitudinal 
ridges; limbs modified to form flippers, 
without claws; marine (Fig. 1A) 
_*Dermochelys coriacea (Linnaeus, 1766) 
Carapace flat, without ridges; feet with 

3 claws; freshwater (Fig. 1B) 
 —— Trionyx euphraticus (Daudin, 1802) 
Limbs modified to form flippers; digits 


not distinct; marine ___.__-------- 4 
Limbs not modified to form flippers; dig- 
its distinct; freshwater or terrestrial __ _ 7 


Four pairs of costal plates on carapace; 
first vertebral plate in contact with mar- 
PINalSnCEIS:, LAP) = ee 5 
Five or more pairs of costal plates on 
carapace; first vertebral plate not in con- 
tact with marginals (Fig. 2C) ________-____- 6 
Plates of carapace imbricate (Fig. 2A) 
__ Eretmochelys imbricata bissa (Ruppell, 
1835) 

Plates of carapace juxtaposed (Fig. 2B) 
___ *Chelonia mydas japonica (Thunberg, 
1787) 

Bridge with 3 inframarginal plates (Fig. 
3A) 
__*Caretta caretta gigas Deraniyagala, 1933 
Bridge with 4 inframarginal plates (Fig. 
3B) 
1829) 

Head covered with undivided smooth 
Skim; digits fully webbed. ..... 2.22..__- 8 
Head covered by shields; digits not 
AMES BL Cee ec Se Na ee neem oR 9 
Axillary and inguinal plates present on 
plastron; plastron united to carapace by 
bony suture; plastron not hinged, im- 
movable; anal plates of plastron pointed, 
their median suture shorter than inter- 
abdominal suture (Fig. 44)... 
ECM aN Mauremys caspica caspica (Gmelin, 
1774) 

Axillary and inguinal plates absent; plas- 
tron united to carapace by ligamentous 
attachment; plastron more or less dis- 
tinctly hinged, movable (in adults); anals 
rounded, their median suture longer than 
interabdominal suture (Fig. 4B) 
ee Emys orbicularis (Linnaeus, 1758) 
Forelimb with 4 claws 
<8, ae Agrionemys horsfieldii (Gray, 1844) 
Forelimb with 5 claws 2... = 10 


*Lepidochelys olivacea (Eschscholtz, 


FIGURE 4. 


505 


(A) Plastron of Mauremys caspica. (B) Plastron 


of Emys orbicularis. Abbreviations: a = axillary, ab = ab- 
dominal, an = anal, f = femoral, g = gular, h = humeral, 


in = inguinal, | = ligamentous attachment, p = pectoral. 


506 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


10a. Shell oval in outline, with smooth, 
rounded posterior margin; ground color 
light olive, with large, distinct, individual 
dank markings Aas s@). hs Pe 
Wien, eke ee Testudo graeca ibera Pallas, 1814 

. Shell elongate in outline, with upturned, 
emarginate posterior margin; ground col- 
or brownish olive, with very indistinct 
darks markings: 4 27 e 2) eA eke 
__ Testudo graeca zarudnyi Nikolsky, 1896 


TESTUDINES 


Family CHELONIIDAE 
Genus Caretta Rafinesque 


Caretta RAFINESQUE, 1814:66 (type-species: Caretta nasuta 
Rafinesque [= Caretta caretta (Linnaeus, 1758)}). 


DEFINITION.—Large marine turtles with five 
or more pairs of costal plates, first in contact 
with nuchal; three poreless inframarginal plates 
on bridge; elongated carapace with vertebral 
keel, becoming smoother with age, highest an- 
terior of bridge, serrated posteriorly (Ernst and 
Barbour 1972:231). 


Caretta caretta (Linnaeus) 
*Caretta caretta gigas Deraniyagala 
Caretta gigas DERANIYAGALA, 1933:66, pl. 5, figs. 4-6 (type- 


locality: Gulf of Mannar, Ceylon). 
Caretta caretta gigas: DERANIYAGALA 1939: 164, figs. 66, 73. 


DIAGNosIs.—Seven to 12 neural bones, last 
one to five usually interrupted by costal bones; 
13 marginal plates on each side of carapace. 

CoLor PATTERN.—Carapace reddish brown, 
sometimes tinged with olive, plates may be bor- 
dered with yellow; plastron yellow to cream. 
Head reddish or yellow-chestnut to olive-brown, 
scales often margined with yellow, and yellow 
below. (Ernst and Barbour 1972:231). 

S1zE.—Averaging about 300 Ibs (136 kg), but 
weights to 1,000 Ibs (455 kg) recorded, with 
lengths up to two meters. 

IRANIAN DISTRIBUTION.—I find no records 
for the Persian Gulf, although it may be expect- 
ed to occur there. 


Genus Chelonia Latreille 


Chelonia LATREILLE, 1801:22 (type-species: Chelonia mydas 
{Linnaeus, 1758)]). 


DEFINITION.—Each flipper with a single claw; 
one pair of prefrontal shields on head; horny 
beak of lower jaw coarsely toothed, that of up- 
per jaw with strong ridges on inner surface; four 


pairs of costal plates, first not in contact with 
nuchal; four inframarginal plates. 


Chelonia mydas (Linnaeus) 

*Chelonia mydas japonica (Thunberg) 

Testudo japonica THUNBERG, 1787:178, pl. 7, fig. 1 (type-lo- 
cality: Japan). 

Chelonia mydas japonica: WERMUTH AND MERTENS 1961:237— 
238. 


DIAGNosIS.—A broad, deep shell, often 
markedly indented above hind limbs. Greenish 
or olive-brown, some individuals melanistic, be- 
coming slate-gray to black. 

SIZE.—Average carapace length of adult 
males about one meter, slightly longer for fe- 
males. 

SEXUAL DIMORPHISM.—Tail of male extends 
well beyond carapace margin, prehensile in ver- 
tical plane; tail of female barely attains edge of 
carapace; the single claw of male long and 
curved. 


IRANIAN LOCALITIES.—Minton (1966:59-61) reports the 
species from the vicinity of Karachi, and Shockly (1949) re- 
ported it nesting at Ras Jawani, Pakistan, close to the Iranian 
border. Although there are no specimens of record, and no 
published reports of green turtles in Iran in the herpetological 
literature, preliminary investigations of the status of marine 
turtles in Iran were carried out by Bosch, Bullock, Kinunen, 
and Walczak. The following sight records were given in their 
reports to the Iran Game and Fish Department in 1970 and 
1971 (Walczak and Kinunen 1970, 1971; Bullock and Kinunen 
1971; Kinunen and Walczak 1971; Walczak 1971). 

Ostan 8 (Fars): 2 km E Bandar-e Maqam [26°56'N, 
§3°29'E]. Ostan 10 (Baluchistan-Sistan): Chah Bahar, 
25°18'N, 60°37'E; 10 km E Chah Bahar; Khalij-e Chah Bahar, 
25°20'N, 60°30’E; 10 km E Pasarvandan [25°05’N, 61°25’E], 
tracks only. Persian Gulf: Jazireh-ye Hormoz, 27°04'N, 
5$6°28'E; Jazireh-ye Larak, 26°52'N, 56°22’E; Jazireh-ye 
Sheykh Sho’eyb, 26°48’N, 53°15'E; Qeshm Island, 26°45'N, 
55°45'E. 


REMARKS.—Minton (1966:59-61) found the 
green turtle nesting primarily from late June 
through early November. Walczak counted 22 
nests attributed to this species 28-29 March 1971 
on Hormoz Island (Walczak 1971). Eggs were 
found 28-30 cm below the sand surface. Al- 
though omnivorous, adults prefer plant material, 
and Bullock and Kinunen (1971) saw green tur- 
tles grazing on algae near shore off Lavan Island 
(Jazireh-ye Sheykh Sho’eyb), usually in groups 
of three to six. They remained submerged to 
feed for five to ten minutes, surfacing only brief- 
ly for air. These authors recorded 52 sightings 
in mid-February (Kinunen and Walczak 1971). 
They found dead females near Chah Bahar and 


ANDERSON: IRANIAN REPTILES 


near Bandar-e Maqam in early May, late Octo- 
ber, and late November. Greatly valued for flesh 
and eggs, populations of green turtles have been 
greatly reduced in many areas of their range. To 
what extent human exploitation has affected 
populations in the Persian Gulf is not known, 
but Kinunen and Walczak (1971) recorded tur- 
tles killed for shells and found nests ravaged by 
people, dogs, and foxes. They found no evi- 
dence of human consumption of turtle flesh, 
proscribed by Muslim religious beliefs. 


Genus Eretmochelys Fitzinger 


Eretmochelys FitZiNGER, 1843:30 (type-species: Eretmoche- 
lys imbricata [Linnaeus, 1766]). 


DEFINITION.—Plates of carapace imbricate 
except in very young individuals; four pairs of 
costal plates; four inframarginal plates; two 
pairs of prefrontal shields on head. 


Eretmochelys imbricata (Linnaeus) 
Eretmochelys imbricata bissa (Ruppell) 

(Figure 5) 

Caretta bissa RUPPELL, 1835:4, pl. 2 (type-locality: Red Sea; 


holotype: Senckenberg Museum no. 7886, 3). 
Eretmochelys imbricata bissa: SMITH AND TAYLOR 1950:16. 


DIAGNosis.—Heart-shaped carapace, fully 
continuous vertebral keel, all vertebrals with 
ridges converging posteriorly; head and flippers 
almost solid black (Ernst and Barbour 1972:223). 

CoLorR PATTERN.—Carapace dark greenish 
brown, plastron yellow. 

SIZE.—To one meter long and about 200 Ibs 
(91 kg). 


IRANIAN LOCALITIES.—T wo skulls from Jabrin are the only 
specimens of record. Kinunen and Walczak (1971) listed sight 
records from four other localities in the Persian Gulf. 

Ostan 10 (Baluchistan—Sistan): Beris, 25°09'N, 61°11'E. 
Persian Gulf: *Jabrin [= Nakhilu Island], 27°51'N, 51°26’E 
(MMTT); Jazireh-ye Hormoz, 27°04'N, 56°28’E; Jazireh-ye 
Sheykh Sho’eyb, 26°48'N, 53°15’E; Jazireh-ye Shotur, 
26°47'N, 53°25’E. 

REMARKS.—These turtles are omnivorous, 
but apparently prefer invertebrates. They share 
nesting beaches with other species of marine 
turtles. Hawksbill turtles are the source of com- 
mercially valuable tortoise shell. There are two 
skulls of this species in the Iranian National 
Museum of Natural History, collected by Derek 
Scott, who said that he had also seen hatchling 
hawksbill turtles, had watched one turtle nest- 
ing, and counted numerous tracks on a small 
island just off the coast in the Persian Gulf. 


507 


Kinunen and Walczak, in unpublished reports 
of their investigations of marine turtles in the 
Persian Gulf, recorded nesting by hawksbills in 
late April and early May and early to mid-June 
1971 on Hormoz Island, on Sheykh Sho’eyb Is- 
land, where they counted 218 nests, and on Shit- 
var Island (Jazireh-ye Shotur), where they found 
250-300 nests on clean white-sand beaches. In 
the vicinity of villages, many nests had been de- 
stroyed by people and dogs. They also found 
remains of these turtles which had been killed 
for the shell. They found three dead turtles on 
the beach at Beris in late October, but all nests 
were found on islands; comparable beaches on 
the adjacent mainland showed no evidence of 
hawksbill nesting. On Chat-e Lambe Beach on 
Hormoz Island, they watched nesting behavior 
of one turtle on 30 April 1971. Their detailed 
account of nesting (Kinunen and Walczak 1971) 
is similar to that described by Carr as cited in 
Ernst and Barbour (1972:225—227). 


Genus Lepidochelys Fitzinger 


Lepidochelys FITZINGER, 1843:30 (type-species: Lepidochelys 
olivacea [Eschscholtz, 1829]). 


DEFINITION.—Four inframarginal plates, at 
least some with pores; first vertebral plate in 
contact with nuchal. 


*Lepidochelys olivacea (Eschscholtz) 


Chelonia olivacea ESCHSCHOLTZ, 1829:2, pl. 3 (type-locality: 
Manila Bay, Philippine Islands). 
Lepidochelys olivacea: FITZINGER 1843:30. 


DiAGNosis.—Usually more than five pairs of 
costal plates; carapace olive. 

CoLor PATTERN.—Carapace Olive, skin olive 
above, lighter below, plastron greenish white or 
greenish yellow. 

S1iz—E.—Carapace usually under 75 cm in 
length; to about 180 Ibs (82 kg). 

SEXUAL DIMORPHISM.—Tail of male thick, 
extending well beyond edge of carapace; tail of 
female usually not reaching margin of carapace; 
plastron of male concave. 

DISTRIBUTION.—No records exist for Persian 
Gulf, but Minton (1966:61) found them breeding 
at Hawke’s Bay, Pakistan. 


Family DERMOCHELYIDAE 
Genus Dermochelys Blainville 


Dermochelys BLAINVILLE, 1816:119 (type-species: Der- 
mochelys coriacea [Linnaeus, 1766]). 


508 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. Ser. 4, Vol. 41, No. 22 


FiGurE 5. Skull of Eretmochelys imbricata bissa, Nakhilu Island, Persian Gulf. MMTT specimen. (Top) Dorsal, (middle) 
ventral, (bottom) lateral. 


ANDERSON: IRANIAN REPTILES 


be * 


ry ts 
f 


DEFINITION.—Carapace covered with smooth, 
leathery skin, and lacking horny plates; cara- 
pace and plastron composed of small platelets 
of bone imbedded in skin, plastral bones other 
than margin absent. 


*Dermochelys coriacea (Linnaeus) 


Testudo coriacea LINNAEUS, 1766:350 (type-locality: Medi- 
terranean Sea). 
Dermochelys coriacea: BOULENGER 1889: 10. 


DiaGNosis.—As for the genus. 

CoLorR PATTERN.—Carapace, head, neck, 
and limbs black or dark brown with small white 
or yellow blotches; plastron whitish. 

SizeE.—Adults to about 2.7 m in length and 
1,600 Ibs (727 kg). 

SEXUAL DIMORPHISM.—Male with concave 
plastron, tail longer than hind limbs; tail of fe- 
male about half as long as hind limbs. 

DISTRIBUTION.—No specimens of record for 
the Persian Gulf, but Minton (1966:59) records 
it from Hawke’s Bay, Pakistan. 

REMARKS.—Kinunen and Walezak (1971) re- 
corded an oil-rendering operation on Jazireh-ye 
Larak, where leatherback turtles are processed 


509 


to provide oil for sealing the hulls of small craft. 
They saw 12 dried skulls there. They did not 
determine the locality at which the turtles were 
obtained, but presumably they were caught in 
the vicinity of that island. 


Family TESTUDINIDAE 
Genus Emys Dumeril 
Emys DUMERIL, 1806:76 (type-species: Emys europaea 


Schweigger, 1814 [= Emys orbicularis (Linnaeus, 1758)], by 
subsequent designation of Fitzinger 1843:29). 


DEFINITION.—Beak mesially notched, not 
hooked; skin on hinder part of head smooth, 
undivided; back of thighs without tubercles; dig- 
its fully webbed; posterior margin of carapace 
smooth; only anterior part of plastron somewhat 
movable, a more or less distinct hinge between 
hyo- and hypoplastra; entoplastron crossed by 
humeropectoral sulcus well in advance of its 
posterior border; plastron united to carapace by 
a ligament. Skull with orbits completely con- 
cealed when viewed dorsally (Loveridge and 
Williams 1957:202). No axillary or inguinal 
plates. 


510 


Emys orbicularis (Linnaeus) 

(Figures 6, 12) 

Testudo orbicularis LINNAEUS, 1758:198 (type-locality: south- 
ern Europe).—PALLAS 1814:17. 

Emys lutaria: STRAUCH 1862:101-—104. 

Cistudo europaea: DE FILippPi 1865:352. 

Emys orbicularis: BLANFORD 1876:308-309. —BOULENGER 
1889:112—114, text-fig. 31. —WERNER 1903:340. —NIKOL- 
SKY 1915:13-25, text-figs. 1, 3-4. —WERNER 1936:200. — 
TERENTJEV AND CHERNOV 1949:107, 323, fig. 33. —For- 
CART 1950:144. —WETTSTEIN 1951:429. —MERTENS 
1957:119. —MERTENS AND WERMUTH 1960:63. —WER- 
MUTH AND MERTENS 1961:79-81, fig. 53 a-<d. —PRITCHARD 
1966:273. —ANDERSON 1968:324—325. —Tuck 1971:52. — 
ANDERSON 1974:30, 42. 


DiaGNosis.—As for the genus (as defined by 
Loveridge and Williams 1957:202). 

COLOR PATTERN.—Carapace light or dark ol- 
ive, dark brown, or black, uniform or with yel- 
lowish dots or radiating lines, or with irregular 
dark markings; plastron dark brown, black, yel- 
low, or brown and yellow; head and limbs dark 
olive, with yellow spots and vermiculations. 

SEXUAL DIMORPHISM.—Loveridge and Wil- 
liams (1957:207) were unable to confirm Boulen- 
ger’s (1889:113) findings of sexual differences in 
tail length and spotting. Pritchard (1966:273) re- 
ported that in two specimens near Astara, Iran, 
the male had red eyes, the female yellowish. He 
did not state how the sex of his specimens was 
determined. 

S1ZE.—Carapace length to 190 mm. 

NATURAL History.—West of Pahlavi, they 
first appear in March, becoming numerous along 
the water courses by April, often climbing up to 
a meter high on logs and brush to bask (Mertens 
1957:119). Aspects of the natural history of this 
species outside of Iran have been discussed by 
Nikolsky (1915:22—24) and by Loveridge and 
Williams (1957:208). 

HaBITAT.—This turtle occurs along the coast 
of the Caspian Sea and has been stated by De 
Filippi to prefer brackish water, such as the 
Murdab, while Mauremys caspica is found in 
the running streams of fresh water (Blanford 
1876:309). Sobolevsky (1929:135) states that in 
the Lenkoran District just north of the Iranian 
border, this species is found in the littoral zone 
of marshes, lakes, and pools of stagnant water 
along the Caspian coast, in the damp subtropical 
forests of the lowlands and lower mountain belt, 
and on the flat, uniform plain of the Moghan 
Steppe. There are two specimens (not examined 
by me) in the Iran National Museum of Natural 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


History from Neur Lake, which is 35 airline km 
east-southeast of Ardabil at 2,400 m elevation. 
This is a spring-fed lake in a small valley near 
the crest of the Talish Mountains. The lake has 
marshy borders and the vegetation of the sur- 
rounding terrain consists of scattered shrubs 
with a “‘pincushion’’ growth form. There are 
large populations of gammarid amphipods and 
aquatic insects in the lake, and in late June there 
are dense aggregations of Rana camerani tad- 
poles in the shallow waters of the lake margins. 
Adults of R. camerani are also extremely nu- 
merous. The only reptiles which I collected in 
this environment were Natrix tessellata and 
Ablepharus bivittatus. A Lacerta, tentatively 
identified as L. saxicola, was seen in the sur- 
rounding hills. At Ardabil, I was given a speci- 
men of Emys orbicularis by the owner of a pub- 
lic bath house, who said he had found it in his 
garden within the city. It may have been an es- 
caped pet. 

ZOOGEOGRAPHIC ELEMENT.—Mediterranean 
(Anderson 1968:332). 


IRANIAN LOCALITIES.—Chartschang, on the Siyahrud (LF 
50); Kura River (AN 15). Ostan 2 (Gilan): Astara, 38°26'N, 
48°52'E (RT 71); 45 km S Astara (RT 71); Bandar-e Pahlavi, 
37°28'N, 49°27'E (WB 76; GB 89; FW 03); Mordab, 37°26'N, 
49°25'E (FdF 65); 20 km E Rasht [37°16’N, 49°36’E] (RT 71); 
Rezvandeh, 37°33'N, 49°09’E, and vic. (RM 57). Ostan 3 (East 
Azarbaijan): * Ardabil, 38°15'N, 48°18’'E (MMTT); Neur Lake, 
38°00'N, 48°34’E, 2,400 m (MMTT). Ostan 12 (Mazanderan): 
*Babol Sar, 36°43’N, 52°39’E (AMNH); *Dasht-e Naz Na- 
tional Park (MMTT); *8 miles N Gorgan [36°50'N, 54°29’E] 
(FMNH); Khorramabad, 36°46’N, 50°53’E (OW 51); Rudkha- 
nehye Talar near Dasteng-Kela (LF S50). 


Genus Mauremys Gray 


Mauremys GRAY, 1870:500 (type-species: Emys fuliginosa 
Gray, 1860, by original designation). 


DEFINITION.—Beak not hooked, skin on hin- 
der part of head smooth, undivided; backs of 
thighs without tubercles; digits fully webbed; 
plastron immovable, united to carapace by a su- 
ture; no plastral hinge; entoplastron crossed by 
humeropectoral sulcus well in advance of its 
posterior border; skull with orbits largely con- 
cealed when viewed dorsally; basioccipital with 
strong lateral tuberosity extending lateral to the 
lagena, forming the floor of recessus scalae tym- 
pani; forearm with numerous transversely en- 
larged scales separated by granular scales 
(Loveridge and Williams 1957:191; McDowell 
1964:245-247). Axillary and inguinal plates pres- 
ent on plastron. The separation of Mauremys 


ANDERSON: IRANIAN REPTILES 


FIGURE 7. 


from the New World Clemmys and the relation- 
ships of the aquatic Testudinidae are discussed 
by McDowell (1964:239-279). 


Mauremys caspica caspica (Gmelin) 
(Figures 7, 13) 


Testudo caspica GMELIN, 1774:59, pls. 10-11 (type-locality: 
Bach Pusahat [= Pirsagat] near Schamachie, Transcauca- 
sia, USSR). 

Clemmys caspica: WAGLER 1830:pl. 24. —BLANFORD 
1876:309-312. —BOULENGER 1889:103. —SIEBENROCK 
1913:177-187, pl. 10, text-figs. 1-8. —NikKoLsky 1915:7-12. 
—WERNER 1917:220; 1936:200. 

Emys caspia: DE Fivipp1 1865:352. 

Clemmys caspica caspica: MERTENS, MULLER, AND RusT 
1934:61. —SCHMIDT 1939:89. —TERENTJEV AND CHERNOV 
1949: 106-107, text-figs. 34, 37. —ForcarT 1950:143. — 
WETTSTEIN 1951:429. —ScCHMIDT 1955:200. —MERTENS 
1957:119, 125. —HELLMICH 1959:2—-3. —MERTENS AND 
WERMUTH 1960:61. —WERMUTH AND MERTENS 1961:59. 
—ANDERSON 1963:434-435, 473. —GUIBE 1966:97. —AN- 
DERSON 1968:325. 

Mauremys caspica: MCDOWELL 1964:239-279, text-figs. 1-4. 

Mauremys caspica caspica: PRITCHARD 
Tuck 1971:52; 1973:13. —ANDERSON 1974:30, 42. 


DIAGNOosiIs.—Posterior margin of carapace 
not serrate; bridge predominantly light, only su- 
ture between marginals dark. 


1966:27 1-272. — 


Syl 


Mauremys caspica caspica, 3 km E Haft Tappeh, Dez River area, Khuzestan, Iran. CAS 141118. 


CoL_or PATTERN.—Plastron dark brown to 
black, each shield with a yellow lateral border; 
bridge yellow with dark sutures; carapace olive 
and olive-brown, with yellowish stripes, forming 
figure-8’s on costals, circles on marginals; lower 
part of marginals yellow with dark sutures, the 
first seven marginals on each side with two black 
spots; head olive, with a yellow stripe on margin 
of snout and under eye; neck with yellow stripes 
bordered by black. 

SEXUAL DIMORPHISM.—Male with slight con- 
cavity on plastron in region of abdominal and 
femoral plates; plastron of female flat or slightly 
convex; anal opening of male more distal than 
in female, beyond posterior margin of carapace 
when tail is extended. 

S1zeE.—Carapace length to 230 mm (Nikolsky 
1915:9). 

NATURAL History.—In suitable bodies of 
water, these turtles occur in considerable num- 
bers, and many can be seen basking together on 
river banks or on floating objects, immediately 
submerging at any hint of danger. According to 
Nikolsky (1915:11) they hibernate, at least in 
Russian territory, from October or November to 
early March. 


512 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


Copulation has been observed in April in the 
Caspian region (Nikolsky 1915:11) and in late 
October in southwestern Iran (Anderson 
1963:434). Egg-laying is recorded for both the 
above areas in June. The eggs measure about 
40 x 30 mm, and the shells have a chinalike tex- 
ture. 

Known dietary items are fishes, worms, tad- 
poles, and small frogs. Individuals have also 
been reported as feeding on ripe melons (Nikol- 
sky 1915:11-12). Undoubtedly they feed on car- 
rion, as my captive specimens readily took strips 
of raw beef. 

Storks are said to feed on the young turtles, 
while vultures (Neophron percnopterus) suc- 
cessfully prey on adults, pecking the flesh from 
beneath the shell (Nikolsky 1915:12). Man may 
take a heavy toll in some areas where turtles 
may be opened to obtain eggs, thought to be 
useful in the treatment of ubiquitous eye ail- 
ments (Anderson 1963:435). 

HABITAT.—This turtle is common in running 
streams, in ponds, and in lakes throughout 
northern and western Iran, in the mountains, 
foothills, and on the western part of the plateau 
wherever water persists throughout the year, or 
at least is present seasonally from year to year 
without fail. I have found these turtles in a great 
variety of habitats throughout western Iran, 
often in cultivated areas where the only water 
was in marshy pastures or small irrigation ditch- 
es. Surrounding vegetation seems to have little 
correlation with their occurrence, although I 
have not found them in heavy forest. Rana ri- 
dibunda usually occurs in the same habitats. The 
greatest concentration of these turtles that I en- 
countered was along the banks of a stream run- 
ning through the village of Marivan (35°31'N, 
46°10’E) in Kordestan, near the Iraq border, 
about 1,200 m elevation. This is a highly eutro- 
phic stream, and the large turtle population is 
probably supported in part by the availability of 
garbage. According to Nikolsky (1915:9), the 
species has been found in the Caspian Sea itself, 
near the mouths of rivers where the salinity is 
low. The same author states that the species has 
been reported from sulphur springs at 40 C. In 
the Lenkoran District, just north of the Persian 
border, Sobolevsky (1929:135) records this tur- 
tle from the littoral zone, stretching along the 
shore of the Caspian, and characterized by 
abundant marshes, lakes, and marshy rice fields. 
It also occurs there in the forests of the lowlands 


and mountain foreland, a zone of damp subtrop- 
ical forest up to 600 or 700 m elevation. He notes 
its presence in the Moghan Steppe as well, a 
flat, uniform plain overgrown by scanty herba- 
ceous vegetation. 

ZOOGEOGRAPHIC ELEMENT.—Mediterranean 
(Anderson 1968:332). The subspecies caspica, 
considered alone, could be regarded as Iranian. 


IRANIAN LOCALITIES.—From below sea level along the 
Caspian Sea to at least 2,000 m in the Zagros Mountains. It 
apparently does not occur east of the Zagros Mountains on 
the Central Plateau. 

Siyahrud, near Reikande (LF 50). Ostan | (Tehran): Arak 
(FS 13). Ostan 2 (Gilan): *Astara, 38°26'N, 48°52’E (USNM; 
RT 71); Bandar-e Pahlavi, 37°28'N, 49°27'E (JG 66); *6 km W 
Chalus [36°38’N, 51°26’E] (FMNH); *10 km W Chalus 
(FMNH); Galigah, 37°30'N, 49°20'E (RM 57); Mordab, 
37°26'N, 49°25'E (WB 76; RM 57); Rasht, 37°16'N, 49°36’E 
(FS 13); Safid Rud, S Rasht (WB 76). Ostan 3 (East Azar- 
batjan): *78 km N Khalkhal on road to Ardabil, 1,720 m 
(CAS). Ostan 4 (West Azarbaijan): *1 km E Gol Tappeh, ca. 
37°50'N, 45°03'E, at margin of Lake Reza’iyeh, dirt road 29 
km N Reza’iyeh, 1,425 m (MMTT); *Mahabad, 36°45'N, 
45°43'E, Lake Shapur I (MMTT); *7 miles NNE Maku 
(39°17'N, 44°31'E] (FMNH); *36°56'N, 46°17’E, 19 km ESE 
Miandow Ab, on road to Shahindezh, 1,350 m (MMTT); *20 
miles N Reza’ iyeh (CAS; FMNH). Ostdn 5 (Kordestan—Ker- 
manshah): Ab-i-Marik, near Namivand [34°22'N, 46°45’E], 20 
miles NW Kermanshah (PP 66); Deh-e Jami, 34°23’N, 
46°14’E, near Karand (PP 66); *Kermanshah, 34°19’N, 
47°04'E (FMNH); E Kermanshah (OW 51); *34°13’N, 
46°04'E, creek near turnoff to microwave station 42 km W 
Kermanshah on road to Shahabad, 1,640 m (MMTT); *8 km 
E Qasr-e Shirin [34°31'N, 45°35’E] (USNM; RT 71); Sah Tang, 
near Karand [34°16’N, 46°15’E] (PP 66); *32 km WNW San- 
andaj [35°19’N, 47°00'E] (USNM; RT 71); *66 km NW San- 
andaj (USNM; RT 71); vic. Sheykhabad, 33°53’N, 48°02’E, 
ca. 30 miles E Kermanshah (PP 66); *Tappeh Sarab, near 
Bijenah, 7 km E Kermanshah (FMNH); Taut Shami, near 
Karand (PP 66). Ostan 6 (Khuzestan—Lorestan): *lake 17 km 
E Haft Gel [31°28'N, 49°30’E] (CAS; SA 63); *32°05’N, 
48°22'E, 3 km E Haft Tappeh, 380 m (CAS); Harmalah, 120 
km NW Ahvaz (WH 59); Istgah-e Dezh, 31°43’N, 48°38’E 
(WH 59); Istgah-e Ezna, 33°27'N, 49°28’E, between Dow Rud 
and Arak (KS 55); *Khorramabad, 33°30'N, 48°20’E (USNM; 
RT 71, 73); *vic. Masjed Soleyman airfield, 31°59'N, 49°15’E 
(CAS; SA 63); Shadegan, 30°40’N, 48°38’E (RM 57); *Shal- 
gahi, 32°15'N, 48°31'E, SE Dezful (FMNH). Ostan 8 (Fars): 
sulphurous springs 15 km NE Borazjan 29°16’N, 51°12’E (KS 
55); Dudej, 29°33’N, 52°59’E, 40 km E Shiraz (OW 51); Fars 
Province (FW 17); *Parishan Lake [= Daryacheh-ye Famur], 
29°30’'N, 51°50'E (MMTT); *Persepolis, 29°57'N, 52°52'E 
(FMNH; KS 39); *5 km SE Pol-e Abgineh [29°33'N, 51°46’E] 
(CAS; FMNH); Rud-e Kor, 29°36’N, 53°18’E, near Persepolis 
(WB 76); Tang-e Karam, 29°06'N, 53°39’E, 70 miles E Shiraz 
(WB 76). Ostan 12 (Mazanderan): *Babol Sar, 36°43'N, 
52°39’E, at mouth of Bahnamir River (MMTT); *near Babol 
Sar (AMNH; *21 km NW Ghala Khash on Kuhsar River, N 
Gonbad-e Kavus [37°17'N, 55°17’E] (CAS; MMTT); Khorra- 
mabad, 36°46'N, 50°53’E (OW 51); Kura River (AN 15); Rud- 
khaneh-ye Talar, near Dasteng-Kela (LF 50); *Shilat Station, 
18 km N by road of Dasht-e Naz National Park (MMTT). 


ANDERSON: IRANIAN REPTILES 


Genus Agrionemys Khozatsky and Mlynarski 

Agrionemys KHOZATSKY AND MVYNARSKI, 1966: 123-125 
(type-species: Testudo horsfieldii Gray, 1844, by mono- 
typy). 


DEFINITION.—Carapace smooth, without 
keels, slightly convex in comparison with most 
testudinines; posterior margin slightly, but dis- 
tinctly serrated. Two suprapygals (metaneurals), 
the second crossed in its median part by the sul- 
cus separating the last vertebral from the caudal. 
Plastral bridge broad, buttresses evenly devel- 
oped; humeropectoral sulcus crosses entoplas- 
tron in its posterior part; no movable elements 
of shell. Concavity of plastron and other male 
sexual features vague or lacking. Forelimbs with 
four digits. (Khozatsky and Mlynarski 1966; 
Mlynarski 1966). 

Khozatsky and Mlynarski have made a case 
for removing Testudo horsfieldii Gray from Tes- 
tudo on the basis of the above characters, 
which, they claim, distinguish it from all other 
living and fossil testudinine genera. 

DISTRIBUTION.—Central Asia east of the Cas- 
pian, and the eastern Iranian Plateau. A single 
species. 

RELATIONSHIPS.—The authors of this genus, 
in comparing it with other testudinine genera, 
both living and fossil, state that it shares taxo- 
nomic characters with African genera as well as 
with Gopherus and the fossil genus Stylemys. 
However, they believe these similarities to be 
convergent and postulate that the combination 
of ‘‘primitive’’ characters (“‘emydoidal”’ ento- 
plastron and shape of suprapygals) and ad- 
vanced specializations (digits of forelimbs re- 
duced to four) indicate that it may be more 
ancient than Testudo, while its relationships to 
other testudinine genera are as yet unclear. 


Agrionemys horsfieldii (Gray) 
(Figures 8A, 12) 


Testudo horsfieldii GRAY, 1844:7 (type-locality: Afghanistan; 
holotype in British Museum). —BLANFORD 1876:308. — 
NIKOLSKY 1897:306. —ZARUDNY 1903:2. —NIKOLSKY 
1915:31-37, text-figs. 6-7. —WERNER 1936:194. —TER- 
ENTJEV AND CHERNOV 1949:109. —MERTENS AND WER- 
MUTH 1960:66. —WERMUTH AND MERTENS 1961:212-213, 
text-figs. 158a—c. —ANDERSON 1963:474; 1968:325; 1974:30, 
42. 

Testudinella horsfieldii: GRAY 1873:13. 

?Testudo ibera: BOULENGER 1889:176-177 (in part; nec Pal- 
las, 1814). 

Testudo baluchiorum ANNANDALE, 1906:75, 205, pl. 2, fig. 1 


513 


FiGureE 8. (A) Agrionemys horsfieldii plastron. (B) Tes- 
tudo graeca ibera plastron. 


(type-locality: Baluchistan; holotype in Indian Museum, 
Calcutta). 

Agrionemys horsfieldii: KHOZATSKY AND M¥YNARSKI 
1966: 123-125. —MVyYNARSKI 1966:219-223, pls. 29-32. — 
OBsT AND MEUSEL 1974:3 ff., figs. 12, 13, 18, col. pl. 6. 


514 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


DIAGNOosIs.—As for the genus. The only 
Southwest Asian tortoise with only four claws 
on the forelimb. 

CoLOR PATTERN.—Carapace yellow or yel- 
lowish brown, each shield with or without a 
black mark, variable in size; black markings on 
plastron may cover almost entire shield, rarely 
absent. 

S1zeE.—Carapace length to 286 mm (Terentjev 
and Chernov 1949:109). 

NATURAL History.—No information is avail- 
able for Iran. Natural history information for 
this species in the USSR is summarized by Ni- 
kolsky (1915:36) and Terentjev and Chernov 
(1949:109). In Pakistan, Minton (1966:66) re- 
cords a few remarks about its habits. 

HaBItTAT.—In the USSR this animal has been 
found on both sandy and loam steppes, moun- 
tains to about 1,300 m, and oases. Minton 
(1966:66) found them in rocky and hilly terrain 
in Pakistan, being most numerous in grassy 
places near springs. In Pakistan they seemed to 
avoid sandy and clay soils, although they occur 
in rocky desert. Minton found them at 1,600- 
2,300 m. They sought refuge in burrows beneath 
boulders or rock outcrops. Niche partitioning 
between this species and Testudo graeca zarud- 
nyi in the area of apparent overlap (Khorasan) 
remains an interesting problem for investigation. 

ZOOGEOGRAPHIC ELEMENT.—Biregional, Ira- 
nian, and Aralo-Caspian (Irano-Turanian). 


IRANIAN LOCALITIES.—Hisomi (FW 36). Ostan 11 (Khor- 
asan): between Akhangerun, 33°25’N, 60°13’E, and Dorokh 
(NZ 03); Kara-ilitschi Mts. (AN 97); Kuhistan [region center- 
ing on 33°N, 57°E] (FW 36); Rud-i-Mil, between Chah-i-Sagak 
and Duruh [32°17'N, 60°30'E] (JG 73; WB 76; GB 89); 13 km 
SW Sarakhs [36°32'N, 61°11’E] on the Turkmen Plateau be- 
side Sarakhs—Mashhad road (MMTT); Tauran (FW 36). Ostan 
12 (Mazanderan): *Almeh, Chondeh ‘Abbas, Mohammad 
Reza Shah National Park (MMTT). 


REMARKS.—The specimens collected near 
Duruh north of Sistan by General Goldschmidt’s 
party were referred by Gray (1873:13) to Tes- 
tudo horsfieldii, and Blanford (1876:308) con- 
curred in this. Boulenger (1889:177) listed these 
with 7. graeca ibera. Blanford points out that 
only the shells have been preserved. 

I have seen only one Persian example of this 
species. I did not encounter this tortoise in the 
field during my travels in Khorasan and Mazan- 
deran. 

Zarudny (1903:2) reported that it was rare in 
Sistan, completely absent to the south, where it 


was replaced by Testudo graeca zarudnyi, and 
common in the vicinity of Bekhars in eastern 
Khorasan. 


Genus Testudo Linnaeus 


Testudo LINNAEUS, 1758:197 (type-species: Testudo graeca 
Linnaeus, 1758, by subsequent designation of Fitzinger 
1843:29). 


DEFINITION.—I follow here the redefinition 
and restriction of the genus by Loveridge and 
Williams (1957:254-255, or perhaps more cor- 
rectly cited as Williams in Loveridge and Wil- 
liams; see footnote, p. 220, ibid.), but I accept 
the removal of 7. horsfieldii Gray by Khozatsky 
and M}lynarski (1966, see above). The following 
characters appear sufficient to separate Testudo 
(sensu stricto) from other testudinine genera 
(Loveridge and Williams 1957:220-—221): Cara- 
pace without hinge; outer side of fourth costal 
subequal to, or smaller than, outer side of third; 
gulars paired; carapace convex, rigid, solid; 
neural bones unreduced; anterior neurals octag- 
onal and quadrilateral; supranasal scales pre- 
sent; nuchal scale present; plastron more or less 
clearly movable in one or both sexes; prootic 
typically concealed dorsally and anteriorly by 
parietal. 

RELATIONSHIPS.—Loveridge and Williams 
(1957:258-260) have outlined their view of re- 
lationships within the genus as they have re- 
stricted and redefined it. In their view, Testudo 
hermanni (southern Europe) is the most primi- 
tive living member of the genus, Testudo graeca 
(Spain, North Africa, southeastern Europe, and 
Southwest Asia) and its close relative 7. mar- 
ginata (Greece) being more specialized. They 
suggest that the dwarf species T. kleinmanni 
(Tripoli, Egypt, and Sinai) may have been de- 
rived from T. graeca. The zoogeographic impli- 
cations of species distributions within the genus 
have been considered briefly by Loveridge and 
Williams (1957:259-260). The availability of fos- 
sil material suggests that a detailed study of the 
relationships among this genus would be prof- 
itable from a zoogeographic standpoint. 


Testudo graeca Linnaeus 

Testudo graeca ibera Pallas 

(Figures 8B, 9, 14) 

Testudo ibera PALLAS, 1814:18-19, pl. 2, figs. 2-3 (type-lo- 
cality: “‘Iberna,’’ region of the central Kura Valley in the 


Caucasus [designated by Mertens 1946:112]). —DE FILipp1 
1865:352. —BLANFORD 1876:306—307 (in part). —NIKOLSKY 


ANDERSON: IRANIAN REPTILES 


’ 


‘ ets yy 

’ a>. 
. > <? 7 ier 7? 
Sweat SY pea ae 


FIGURE 9. 


1907:260; 1915:25-31. —WERNER 1936:200. —ROSTOMBE- 
KOV 1938:11. 

Testudo graeca: PALLAS 1814:17-18, pl. 2, fig. 1. —TER- 
ENTJEV AND CHERNOV 1949:108-109, 298. —FORCART 
1950:144. —ScHmiptT 1955:200. —PRITCHARD 1966:273- 
274. 

Testudo ecaudata PALLAS, 1814:19, pl. 3, figs. 1-2 (type-lo- 
cality: forest region of Persia on the Caspian Sea). 

Testudo buxtoni BOULENGER, 1920:251 (type-locality: Manjil, 
between Rasht and Kasvin, on south coast of Caspian Sea; 
holotype in British Museum). 

Testudo graeca ibera: MERTENS 1946:112. —WETTSTEIN 
1951:429-430. —MERTENS 1957:120. —MERTENS AND 
WERMUTH 1960:65. —WERMUTH AND MERTENS 1961:208- 
209, text-figs. 156a—d. —ANDERSON 1963:474. —GUIBE 
1966:97. —ANDERSON 1968:325. —TuUck 1971:53. —OBsT 
AND MEuSEL 1974:3 ff., figs. 5, 9, col. pls. 3-4. —ANDER- 
SON 1974:30, 42. 

Testudo zarudnyi: SCHMIDT 1939:89 (nec Nikolsky, 1896). 
—WETTSTEIN 1951:430. —Tuck 1971:53, fig. 2. 

Testudo graeca zarudnyi: PRITCHARD 1966:274 (in part; nec 
Nikolsky, 1896). 


Testudo graeca ibera, 22 km E Ahar, East Azarbaijan, Iran. MMTT specimen. 


DrAGNosis.—Maxillary with a weak ridge; 
quadrate enclosing stapes; forelimb with five 
claws; a large subconical tubercle on posterior 
aspect of thigh; supracaudal shield undivided; 
fifth vertebral shield not broader than third; shell 
oval in outline, with smooth, rounded posterior 
margin (but see remarks below under 7. g. za- 
rudnyi). 

CoLor PATTERN.—Carapace light olive or 
yellowish; scutes with individual black spots, 
larger and less regular in adults than in young, 
sutures sometimes black-margined; plastron 
with dark spots which often nearly cover scutes. 

S1zeE.—Nikolsky (1915:28) gives the maxi- 
mum carapace length as 350 mm. Most speci- 
mens are under 300 mm, however. 

NATURAL History.—Pritchard (1966:273-274) 
observed copulation in this species in the Zagros 


516 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


in late August and early September, whereas 
Nikolsky (1915:30) records mating in April and 
May in the Transcaucasian area. Pritchard has 
described the copulatory behavior observed in 
Iran. Further notes on the natural history of this 
tortoise in areas outside of Iran are provided by 
Nikolsky (1915:31), Terentjev and Chernov 
(1949:108-109), and Reed and Marx (1959:115- 
116). 

HABITAT.—These tortoises are reported from 
elevations near the shore of the Caspian Sea to 
high mountainous localities. They occur on dry 
open steppes, hillsides, and mountains as well 
as cultivated regions and forest situations (Ni- 
kolsky 1915:30; Sobolevsky 1929:137). Rostom- 
bekov (1938:11) records it from Mt. Sitaver at 
about 4,000 m, an elevation which I feel requires 
confirmation for this species. I collected this 
species in dry cultivated areas, in overgrazed 
steppe vegetation on rocky alluvium, on rocky 
hillsides in scrub-oak vegetation, and in grassy 
areas in the vicinity of streams and ponds, up to 
2,050 m elevation. Reed and Marx (1959:115- 
116) discuss the habitat in Iraq. 

DISTRIBUTION.—In northern and western 
Iran, it occurs in AZarbaijan, the Caspian coast, 
and presumably, the Alborz Mountains. The 
subspecific status of the tortoises of the Zagros 
Mountains remains unclear (see below under T. 
g. zarudnyi), but those records are included here 
under 7. g. ibera. From below sea level to above 
2,000 m, possibly to 4,000 m in Iran. 

ZOOGEOGRAPHIC ELEMENT.—The species as 
a whole is biregional, Mediterranean and Ira- 
nian. The subspecies 7. g. ibera is interpreted 
here as Iranian. Wermuth and Mertens (1961:209) 
recognize four subspecies of Testudo graeca: T. 
g. graeca Linnaeus (southern Spain and North 
Africa from Morocco to Cyrenaica), T. g. ibera, 
T. g. terrestris Forskal (Syria and Israel to 
northeastern Africa from middle Egypt west to 
Libya; I know of no records for North Africa 
which can be referred to this subspecies, and 
Loveridge and Williams [1957] and Marx [1968] 
record only 7. kleinmanni Lortet from Egypt 
and Sinai), and 7. g. zarudnyi (see below). Ex- 
cept that its tolerance of aridity enables the sub- 
species 7. g. zarudnyi to occupy the Central 
Plateau in Iran, the distribution of Testudo grae- 
ca Closely parallels that of Mauremys caspica. 
It is noteworthy, however, that no Testudo 
seems to have been collected in the Arabian 


Peninsula proper, while M. caspica occurs at 
least along the Persian Gulf coast of Arabia. 


IRANIAN LOCALITIES.—Forest region on Caspian Sea (PSP 
14, type-locality for T. ecaudata Pallas); Mt. Sitaver, Kor- 
destan 12,000 ft (VR 38). Ostan I (Tehran): W Arak, [34°05'N, 
49°41'E] (OW 51); *Hesarak, 35°51'N, 50°56’E (CAS). Ostan 
2 (Gilan): Bandar-e Pahlavi, 37°28’N, 49°27’E (RT 71); *2 
miles N Bijar [35°52'N, 47°36’E] (FMNH); coastal region of 
SW Caspian coast (RM 57); *1 mile S Divandarreh [35°55'N, 
47°02'E] (FMNH); Manjil, 36°44’N, 49°24’E, between Rasht 
and Qazvin, 7,000—7,500 ft (GB 21, type-locality for T. buxtoni 
Boulenger); *87 miles N Zanjan [36°40'N, 48°29’E] (FMNH). 
Ostan 3 (East Azarbatjan): *38°26'N, 47°15'E, 22 km E Ahar, 
on road from Meshkin Shahr, 1,800 m (MMTT); *37°22'N, 
46°30'E, SE Shalivan, paleo site 29 km E Maragheh, 1,800 m 
(MMTT); *9 km E Maragheh [37°23'N, 46°13'E] (MMTT). 
Ostan 4 West Azarbatjan): *23 miles SSE Reza’iyeh 
[37°33'N, 45°04'E] (FMNH); *near Daryacheh-ye Reza’iyeh 
(MMTT). Ostan 5 (Kordestan—Kermanshah): Bisotin, 34°23'N, 
47°26'E, near Kermanshah (OW 51); 17.5 km SW Borujerd 
[33°54’N, 48°46’E] (RT 71); *21 km N Hoseynabad [35°33'N, 
47°08'E] on road from Sanandaj to Saqgez, 2,050 m (MMTT); 
*34°29'N, 46°41’E, 31 km NW Kermanshah on road to Now- 
sud, 1,440 m (MMTT); *34°13’N, 46°41’E, creek near turnoff 
to microwave station 42 km W Kermanshah on road to Sha- 
habad, 1,640 m (MMTT); 24 km W Karand [34°16'N, 46°15’ E] 
(RT 71); 42 km W Kermanshah [34°19'N, 47°04'E] (RT 71); W 
Kermanshah (OW 51); *3 miles W Lalabad and ca. 25 miles 
NW Kermanshah (FMNH); 32 km WNW Sanandaj [35°19’N, 
47°00'E] (RT 71). Ostan 6 (Khazestan—Lorestan): Bakhtiari 
region (AN 07; 15); Dow Rud, 33°28’N, 49°04’E, 1,400 m (KS 
55); 5 km NW Dow Rud (RT 71); *Khorramabad, 33°30'N, 
48°20'E (FMNH). Ostan 7 (Esfahan): W Esfahan [32°40'N, 
51°38’E] (OW 51); *Galatappeh, 33°13'N, 51°45'E (FMNH); 
15 miles E Murcheh Khvort [33°06’N, 51°30’E], 33 miles N 
Esfahan (PP 66). Ostan 8 (Fars): Khaneh Kowreh, 30°52'N, 
53°09’'E, SE Abadeh, 2,180 m (OW 51); *Yazd-e Khvast, 
31°31'N, 52°07'E (FMNH; KS 39). 


Testudo graeca zarudnyi Nikolsky 
(Figures 10, 14) 


Testudo ibera: BLANFORD 1876:306-307 (in part; nec Pallas, 
1814). —WERNER 1895:1. 

Testudo zarudnyi NIKOLSKY, 1896:369 (type-locality: moun- 
tains in Birjand Province, Iran [restricted by Mertens 
1946:113]; holotype: Zoological Institute, Leningrad, no. 
8738); 1897:307, pl. 17; 1899:375-376. —ZARUDNY 1903:2. 
—WERNER 1929:238-239; 1936:194-195; 1938:265-267, 
text-fig. 1. —ForcarT 1950:144. —ANDERSON 1963:474. 

Testudo graeca zarudnyi: MERTENS 1946:113; 1956:91. — 
WERMUTH AND MERTENS 1961:210. —PRITCHARD 1966:274. 
—ANDERSON 1968:325. —OBsT AND MEUSEL 1974:5 ff. — 
ANDERSON 1974:30, 42. 


DraGnosis.—Maxillary with a weak ridge; 
quadrate enclosing stapes; forelimb with five 
claws; a large subconical tubercle on posterior 
aspect of thigh; supracaudal shield undivided; 
fifth vertebral shield not broader than third; shell 


ANDERSON: IRANIAN REPTILES 


FiGure 10. Testudo graeca zarudnyi, °, 


elongate in outline, with upturned, emarginate 
posterior border. 

CoLor PATTERN.—Carapace brownish olive, 
with very indistinct dark markings. 

S1zE.—To 275 mm carapace length. 

REPRODUCTION.—An adult male in captivity 
almost daily attempted copulation with four fe- 
males during May, June, and July. He would 
ram a female with the front of his plastron and 
then mount without further ado, inverting his 
tail beneath her carapace and opening his mouth 
during the process. The females seemed oblivi- 
ous to these attentions, continuing to eat or to 
walk slowly. I could not tell how frequently he 
was successful in his attempts. He seemed tire- 
less in his efforts, and it seems unlikely he would 
ever encounter such frequent opportunity in his 
natural habitat. 


32 km W Zahedan, Baluchistan, Iran. MMTT specimen. 


Mr. Reza Khazai’e of the Iran National Mu- 
seum of Natural History and I found a nesting 
female at 1000 hrs on 18 April 1975 in rocky hills 
west of Zahedan. She had excavated her nest on 
a west-facing slope, under and between two 
small Artemisia bushes. The nest was 11 cm 
deep, 15 cm long, and 12 cm wide. When locat- 
ed, the tortoise had already laid three eggs, and 
she laid a fourth as we watched. Her activities 
did not seem to be influenced by our presence. 
She began the laborious process of filling the 
nest by scraping in dirt with her hind limbs. Dur- 
ing the hour required for this filling, she did not 
move her body off the excavation, but stretched 
out her hind limbs to the maximum, even at right 
angles to the body, in order to rake in dirt. She 
filled from back to front, resting briefly now and 
then. She scraped the silty alluvium into a fine 


518 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


powder as she worked, and rejected stones, 
kicking them out. She moved forward across the 
nest as she finished, making no obvious tamping 
movements, and raked dead branches and twigs 
over the nest, again with her hind limbs only. 
She did not urinate on the nest, although she 
might have passed water into the excavation 
prior to laying. She moved off the nest and ap- 
peared weak at first as she walked away toward 
the top of the slope. We uncovered the eggs, 
which had 9 cm of earth covering them. When 
we showed the tortoise the exposed nest, she 
seemed interested, bobbing her head, and head- 
ed right for it, covering it with her body. She 
made no attempt to fill the nest again, and may 
simply have been trying to excape from us under 
the bush. The soil at the bottom of the nest was 
slightly damp. Mr. Khazai’e’s subsequent at- 
tempt to incubate the eggs in the laboratory was 
unsuccessful. 

HaBitTAT.—Blanford (1876:306—-307) states 
that in the southern part of Iran, tortoises live 
on barren hillsides and plains far from cultiva- 
tion. The two tortoises I collected in Balichistan 
were in rocky terrain with steep slopes, barren 
except for widely scattered small shrubs. One 
individual had a maggot-infected wound on the 
tail. The shell was scarred and broken, appar- 
ently from falls, which must be frequent in this 
terrain. Perhaps worthy of note is that an adult 
T. g. ibera fell three stories from a balcony in 
Tehran onto concrete without sustaining appar- 
ent injury. Four months later it was still alive 
and seemed in good health. 

DISTRIBUTION.—Iran, on the Central Plateau. 
1,000-—3,000 m elevation. While they have been 
taken close to the eastern border of Iran, these 
tortoises are as yet unknown from Pakistan, 
Afghanistan, and Sistan, where the only recorded 
tortoise is Agrionemys horsfieldii. 

ZOOGEOGRAPHIC ELEMENT.—Iranian; en- 
demic to the Central Plateau. 

IRANIAN LOCALITIES.—Askalabad, Baltchistan, 1,800 m 
(FW 38). Ostan 7 (Esfahan): ‘Arisan, 33°29'N, 54°56’E, in 
the S of the Dasht-e Kavir, 928 and 1,065 m (FW 29; 36). 
Ostan 9 (Kerman): Kerman, 30°17'N, 57°05’E, 6,000 ft (WB 
76; GB 89); vic. Kerman (FW 95); Kuh-e Jupar, 29°55’N, 
57°15'E, ca. 2,700 m (FW 95); steppe between Nabid [29°40’/N, 
57°38'E] and Bam [29°06'N, 58°21’E] (LF 50). Ostan 10 (Ba- 
luchistan—Sistan): Bendun, 31°23'N, 60°44'E (AN 99; NZ 03); 
Gurmukh (NZ 03); *ca. 27°54'N, 60°48’E, 93 km N Iranshahr 
on road to Khash, 1,350 m (MMTT); Sangun, 28°35'N, 


61°20’E, E Kth-e Taftan, 1,700 m (RM 56); Tagab, 17 km SE 
Bazman [27°49'N, 60°12'E] (NZ 03); *ca. 29°28'N, 60°41’E, 


32 km W Zahedan on road to Chashme Ziarat, 1,900 m 
(MMTT). Ostan 11 (Khorasan): Birjand, 32°53'N, 59°13'E 
(AN 96; 97; type-locality); Fandokht, 33°47’N, 59°54’E, Zirk- 
uch (AN 99); Kharestan, 32°17'N, 59°56’E, Sarhaad, 2,500 m 
(FW 38); Kuh-i-Akhangerun, ca. 33°25'N, 60°13’E (NZ 03); 
Kuh-i-Atkul (NZ 03); Kuh-i-Khadzi (NZ 03); Kuh-i-Ma- 
gomed-abad (NZ 03); Sarhad-e Bala (FW 36); Tscha-i- 
Gjuische (AN 99). 


REMARKS.—The taxonomic status of the tor- 
toises of the Central Plateau and the Zagros 
Mountains remains problematical. Blanford 
(1876:307) at first regarded his specimens from 
Kerman as distinct from ibera and stated that 
others examined from Shiraz and on the road 
from Shiraz to Esfahan did not differ from those 
collected at Kerman. After examining tortoises 
from various parts of southern Europe and 
northern Africa, he decided that there were no 
constant differences between these populations 
in the characters (mostly proportions of plastral 
shields) which he considered. Nikolsky 
(1896:369) believed them distinct in the shape of 
the carapace and in color pattern, and additional 
specimens collected by Zarudny in 1898 (Nikol- 
sky 1899:375—376) strengthened his view. Bird 
(1936:259) examined a paratype in the British 
Museum and questioned the separation of za- 
rudnyi even at the subspecific level, while Prit- 
chard (1966:274) examined a paratype also 
(doubtless the same specimen seen by Bird) and 
voiced similar doubts. Schmidt (1939:89) and 
Wettstein (1951:430) identified specimens from 
the Zagros region as zarudnyi. 

I have had the opportunity to examine only 
three specimens from the eastern part of the 
Central Plateau, and have seen only a few spec- 
imens of undoubted ibera from Iran, so I can 
offer only speculation about the status of these 
taxa. I have seen a number of specimens from 
the Zagros region, including Schmidt’s series 
from Yazd-e Khvast. I am impressed with the 
variation in these tortoises, even in the relatively 
small sample I have examined. In most of the 
Zagros specimens, including those from Iraq, 
the dark markings of the carapace are indistinct, 
the shells being almost a uniform dark brown; 
in several, the corners of the shell flare outward 
somewhat, and the posterior margin is emargin- 
ate. In all specimens I examined, the greatest 
height of the carapace is at the third vertebral 
plate; in the specimens from Yazd-e Khvast, the 
profile is somewhat different from most other 
specimens in that the height of the shell at the 


ANDERSON: IRANIAN REPTILES 


fourth vertebral is nearly as great as at the third, 
whereas in most specimens of ibera and graeca 
I examined, the fourth is distinctly lower. The 
profile of the smallest of four specimens from 
Khorramabad is similar to these Yazd-e Khvast 
animals. In the flaring of the shell, FMNH 
21028, from Yazd-e Khvast, probably a female, 
is the extreme; all of the posterior marginals 
flared outward, forming a pronounced ‘“‘skirt.”’ 
This specimen bears a close resemblance in the 
shape of the shell to the specimen figured by 
Nikolsky (1897:pl. 17). Of the specimens I ex- 
amined, the one closest to it in this regard is 
probably FMNH 74950 from Iraq. Specimens 
from Azarbaijan collected by the Street Expe- 
dition to Iran in 1962 also have somewhat flar- 
ing, emarginate shells and indistinct markings, 
while young specimens collected by the 1968 
Street Expedition in both Kordestan and Azar- 
baijan have the shells much less flared, the dark 
spots more individual, many of the plates with 
one Or more margins outlined with dark brown, 
contrasting sharply with the horn color of the 
background. 

Both Robert Tuck of the Iran National Mu- 
seum of Natural History, and Jeromie Anderson 
of Quetta (in litt.) have called my attention to 
specimens from the eastern central Zagros 
which exhibit characteristics attributed to T. g. 
zarudnyi. On the basis of the limited material I 
have examined, I find no consistent, geograph- 
ically correlated distinctions which would justify 
separating the Zagros tortoises from T. g. ibera, 
and the Zagros localities are listed under that 
form. Since recording the above observations, 
I collected two specimens of 7. graeca in east- 
ern Iran and about a half-dozen in northwestern 
Iran. I did not find the opportunity to make care- 
ful comparisons and measurements of these 
specimens, which are still in Tehran, nor have 
they been compared as yet with any of the ma- 
terial mentioned above. The animals from Ba- 
luchistan were distinguishable from those from 
Azarbaijan on the basis of their more flared car- 
apace margins. A male 7. g. zarudnyi seemed 
to make no distinction between the females from 
Balichistan and those from Azarbaijan in his 
amorous attentions. 

I tentatively recognize the taxon T. g. zarud- 
nyi as Valid for tortoises from the eastern Central 
Plateau. It seems likely that clinal or ecotypic 
Variation will be shown to exist, linking the nom- 


519 


inal taxa of Persian tortoises. Should the eastern 
animals prove to be consistently distinct from 
ibera, the Zagros populations may represent a 
broad zone of intergradation between these sub- 
species. 

In many Iranian specimens the first claw of 
the forelimb is quite small, and if overlooked, 
could prove misleading in use of the key. 


Family TRIONYCHIDAE 

Genus Trionyx Geoffroy 
Trionyx GEOFFROY, 1809:4 (type-species: Trionyx aegyptia- 
cus Geoffroy, 1809 [= Trionyx triunguis (Forskal, 1775)] by 
original designation; see Schmidt 1953:108; Loveridge and 


Williams 1957:422-423; Wermuth and Mertens 1961:xxii- 
Xxilil, 260). 


DEFINITION.—Plastron without cutaneous 
femoral flaps; skull without maxillary ridging; 
orbit nearer the temporal than the nasal fossa; 
intermaxillary foramen moderate to large; pre- 
frontal always in contact with vomer; jugal not 
or but scarcely in contact with parietal; postor- 
bital arch narrower than diameter of orbit; pro- 
boscis as long as eye opening (Smith 1931:154; 
Loveridge and Williams 1957:423). 

RELATIONSHIPS.—Of the living genera of 
trionychids, Trionyx is the least specialized, the 
most widely (but disjunctly) distributed, has the 
longest fossil record, and the greatest number of 
species. Loveridge and Williams (1957:416) pre- 
sent a dendrogram summarizing their interpre- 
tation of the relationships within the genus. 
They regard the monotypic genera Pelochelys 
and Chitra as specialized offshoots of the Ori- 
ental subgroup of Trionyx. 


Trionyx euphraticus (Daudin) 

(Figure 12) 

Testudo euphraticus DAUDIN, 1802:305 (type-locality: banks 
of the Euphrates River). 

Trionyx euphraticus: GEOFFROY 1809:17. —BLANFORD 
1876:312-313. —ANDERSON 1963:474; 1968:325; 1974:30, 
42. 

Amyda euphratica: MERTENS 1957:125—126. 


DIAGNosIs.—Eight pairs of pleural bones; 8th 
pleurals partially reduced; young with numerous 
longitudinal ridges of dermal tubercles; carapace 
without ocelli. 

COLOR PATTERN.—Carapace uniform dark 
green in adults, venter white; young specimens 
have large light spots on top of head, numerous 
smaller light flecks on carapace. 


520 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


S1zeE.—The largest recorded Iranian specimen 
has a carapace length of 290 mm (Mertens 
1957:125). Boulenger (1889:258) gives 370 mm 
as the size of a specimen from the Euphrates. 

NATURAL History.—These turtles are very 
wary, disappearing into the water when ap- 
proached. According to Richter, they can be 
taken readily with hooks. When captured, they 
bite viciously. (Mertens 1957:126) 

I have found little published information on 
the natural history of this species. However, 
soft-shells generally are known to be rather thor- 
oughly aquatic, but able to move rapidly on land 
as well as in the water. They inhabit large bodies 
of permanant water and shun temporary streams 
and ponds. They are known for their habit of 
burying themselves in the sand or mud of stream 
and lake bottoms. They bask on banks and 
shores, but never far from the water. These tur- 
tles are actively predacious, feeding on fish, 
frogs, crustaceans, mollusks, and insect larvae; 
vegetable matter is consumed by at least some 
species. Webb (1962:541-577) summarizes the 
literature on the American species of Trionyx, 
and the habits of Old World species are doubt- 
less similar. 

RELATIONSHIPS.—According to Loveridge 
and Williams (1957:416-417) the closest living 
relative of Trionyx euphraticus is T. swinhoei. 
These two species, which have reduced eighth 
pleurals, are more closely related to New World 
species than to other Old World species, ac- 
cording to these authors. However, Webb 
(1962:579-580) states that the American 7. ferox 
is more closely related to Old World species 
than to the other three New World species. He 
does not say to which Old World species 7. fer- 
ox is related, but mentions the large eighth pair 
of pleurals as one character separating it from 
other American soft-shells. 

DISTRIBUTION.—The Tigris and Euphrates 
drainages; known from southern Turkey, north- 
eastern Israel, Syria, Iraq, and Iran. In Iran, 
specimens have been collected at a single local- 
ity in Khizetan, but species probably occurs 
in much of the Karun River drainage, as the 
Karun empties into the Shatt-al-Arab, the con- 
fluence of the Tigris and Euphrates. 

ZOOGEOGRAPHIC ELEMENT.—Difficult to as- 
sign with certainty, as the disrupted distribution 
of the genus as a whole represents the fragmen- 
tation of a once more widespread range. I regard 
T. euphraticus as Saharo-Sindian (Anderson 


1968:332), while Bodenheimer (1944:71) consid- 
ered it Irano-Turanian. 


IRANIAN LOCALITIES.—Ostan 6 (Khuzestan—Lorestan): 
Shadegan, 30°40’N, 48°38’E, on the Rudkhaneh-ye Jarrahi 
(RM 57). 


REMARKS.—Recently, a specimen was cap- 
tured, photographed, and released in the Karun 
River (Tuck, in litt.). 


CROCODILIA 


Family CROCODYLIDAE 
Genus Crocodylus Laurenti 


Crocodylus LAURENTI, 1768:53 (type-species: Crocodylus nil- 
oticus Laurenti, 1768). 


DEFINITION.—*‘ 16-19 upper and 14-15 lower 
teeth on each side; fourth or fifth maxillary tooth 
largest; fourth mandibular tooth fitting into a 
notch in the upper jaw. Snout more or less elon- 
gate; nasal bones extending to the nasal opening; 
splenial bones not entering the mandibular sym- 
physis, which does not extend beyond the eighth 
tooth. A dorsal shield formed of from four to 
eight longitudinal series of keeled bony scutes.” 
(Smith 1931:40) 


Crocodylus palustris Lesson 

(Figure 12) 

Crocodilus palustris LESSON, 1834:305 (type-locality: Ganges 
River). 

Crocodylus palustris palustris: DERANIYAGALA 1936:282. — 
HONEGGER 1971:45-46. 


DIAGNosIs.—Nineteen teeth in upper jaw on 
each side, 5 in each premaxillary; premaxillo- 
maxillary suture on palate nearly straight across; 
width of snout about 60 percent of length, usu- 
ally without distinct ridges in front of eye; 4 or 
5 large nuchal plates; 4 postoccipital plates in a 
transverse series; dorsal plates in 16-18 trans- 
verse, and 4 or 6 longitudinal series; 16 rectan- 
gular smooth plates across middle of belly. 
(Smith 1931:47; Minton 1966:71) 

CoLor PATTERN.—Brown or olive above with 
dark mottling to almost uniform black, yellowish 
below; young lighter, olive with dark blotches. 

S1zE.—Smith (1931:47) gives four meters as 
maximum size, but Minton (1966:71) stated that 
the largest individuals he saw in Pakistan were 
about three meters total length. The largest 
crocodiles observed by biologists in Iran have 
been estimated at no more than two meters total 
length. 


ANDERSON: IRANIAN REPTILES 


NATURAL History.—Little seems to be 
known of the habits of the marsh crocodile, or 
at least there is little information that is reliable 
in the herpetological literature (Neill 1971:416). 
Smith (1931:48) and Minton (1966:71) have sum- 
marized what little is known of the behavior of 
this animal in India and Pakistan. Smith says 
that it only occasionally attacks man, and Min- 
ton says that he knows of no authenticated ac- 
counts of attacks on humans in Pakistan. Neill 
states that there is no basis for stories that these 
crocodiles eat living people or corpses. Insofar 
as I could determine, villagers do not fear them 
in Baluchistan (where many harmless animals 
are feared), and children swim in pools claimed 
to be inhabited by crocodiles. In Iran, these an- 
imals are shy and difficult to observe. Appar- 
ently, they are not considered a hazard to do- 
mestic stock in Baluchistan. 

The only recorded observations of crocodiles 
in Iran are those of Kinunen and Bullock (1970) 
and Bosch, Bullock, and Kinunen (1970), in un- 
published reports of the Iran Division of Re- 
search and Development. They reported croc- 
odile tracks entering and leaving a ‘“‘den’’ about 
two meters above water level excavated in a 
sloping bank. The entrance was 80 cm wide and 
40 cm high. The excavation was at least two 
meters deep, enlarged into an inner chamber 
lower than the entrance. During the period of 
observation by these investigators (mid-Novem- 
ber), crocodiles spent much of the day basking 
on the river banks or floating and swimming. 
The report states that at this time of year the 
crocodiles were able to remain submerged for at 
least four or five hours. 

Nothing is known of the reproduction in Iran. 

Minton (1966:71) says that fish and turtles 
seem to be their principal food in Pakistan. Neill 
(1971:416) says that their food includes fishes, 
frogs, and sizable aquatic mammals, with lesser 
numbers of smaller mammals and birds. No 
aquatic turtles are known from Baltchistan, nor 
are there any aquatic mammals. Possible croc- 
odile food items in this region include cyprinid 
fishes, and in the lower reaches of the Sarbaz 
River, mud-skippers (Periophthalmus sp.). 
Frogs (Rana cyanophlyctis) occur in this area. 
Small mammals, including palm squirrels, 
groundsquirrels, foxes, mongooses, and gazelles 
may fall victim occasionally to crocodiles when 
they visit the pools in the river to drink. Water 
birds are available in this area, and Bosch, Bul- 


521 


lock, and Kinunen observed a crocodile near a 
dead and mutilated mallard duck on a river 
bank. 

Young crocodiles likely fall prey to the larger 
mammalian and avian predators, and possibly to 
adults of their own species. Desert monitors 
(Varanus griseus) are possible egg predators, as 
may be foxes, mongooses, and domestic dogs. 
Man is the only predator on adults, and hide 
poachers pose a continuing threat to the exis- 
tence of the species in Balichistan. Kinunen and 
Bullock (1970) state that Pakistani hide-hunters 
were reported to have taken 18 crocodiles from 
the Sarbaz River between Rask and Bahu Kalat 
in 1967. 

HaBItAt.—Throughout its range, the marsh 
crocodile inhabits freshwater marshes, pools, 
and rivers. Probably the only suitable crocodile 
habitat in Iranian Balichistan is along the Sar- 
baz River, although the Makran coast between 
Jask and Chah Bahar should be carefully ex- 
plored for other possible crocodile localities. 
The valuable report of Bosch, Bullock, and Kin- 
unen describes the habitat of those areas in 
which they saw crocodiles. The headwaters of 
the Sarbaz River lie in the mountains of the Per- 
sian Makran. The river runs through a narrow 
canyon before forming a broad floodplain below 
Garm-e Bit. This floodplain is several kilometers 
wide and opens into Gavater Bay of the Gulf of 
Oman west of Jiwani. The flow of the river is 
subject to great seasonal fluctuation and yearly 
variation, being intermittent during the dry sea- 
son and subject to flooding during the rainy sea- 
son. The climate of the region is hot and dry, 
annual rainfall being less than 10 cm. Dominant 
vegetation along the river is Tamarix, Acacia, 
and Prosopis, date palms being cultivated 
around villages. During the dry season, only 
pools up to two meters in depth and a kilometer 
in length persist in much of the lower reaches of 
the river, and a significant portion of the flow 
below Rask is used for agricultural purposes. 
During flood times, the entire river bed may be 
flooded to three or four meters above normal 
water level. Bosch, Bullock, and Kinunen (1970) 
observed crocodiles in pools at least 1.5 m deep, 
under 10—20 m high banks, usually at a bend of 
the river or oxbow pools. Crocodiles were never 
seen in pools in the middle of the valley. Usually 
there were shrubs on the banks, and often reeds 
were present. At Kolani the river is 10-20 m 
wide, running continuously for several kilome- 


$99 


22 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


FiGure 11. 


ters. It is bordered by a thick growth of reeds, 
shrubs, and trees. I observed only one croco- 
dile, near the village of Hudar. The river bed 
here is lined with Tamarix and oleanders (Ner- 
ium). Date orchards in the floodplain support the 
village, which sits on a slope above the river. 
The pool, through which the river was flowing 
in late April, was near the base of a bluff. Tam- 
arisks and oleanders overhung the east bank of 
the pool. The water was fairly turbid. We had 
found the spot where crocodiles had basked on 
the river bank, and had seen tracks, but the only 
animal seen was floating in the water near the 
far bank. It was a relatively small individual, 
about 1.5 m total length. We could get no closer 
than about 25 or 30 m. 

DIsTRIBUTION.—Assam west through most of 
India and Sri Lanka, intermittently through Ba- 
lichistan to extreme southeastern Iran. In Pak- 
istan it occurs throughout the Indus Valley north 
at least to Multar and Bahawalpur; the Hab Riv- 
er at least to Diwana. On the Makran coast in 
the Hingol and Dasht Rivers. In Iran, it is known 
only from the Sarbaz River, the drainage just to 
the west of the Dasht River of Pakistan. The 
crocodiles of Sri Lanka are considered a distinct 


Diplometopon zarudnyi, Dhahran, Saudi Arabia. CAS specimen. 


subspecies, C. p. brevirostris (Werner). At pre- 

sent, all other populations, including those from 

Iran, are presumed to be C. p. palustris. 
ZOOGEOGRAPHIC ELEMENT.—Indian. 


IRANIAN LOCALITIES.—AIlI of the following localities are 
sight records, the one from Hudar my own, the others those 
of Bosch, Bullock, and Kinunen. There are no specimens of 
record at present. Ostan 10 (Balichistan—Sistan): Baht Kalat, 
25°43'N, 61°25’E; 25°55'N, 61°34’E, 30 km N Bahu Kalat, 150 
m; Djork-e Bit, road construction camp; 3 km below Firuza- 
bad [26°15’N, 61°23'E]; Garm-e Bit; 4 km above Garm-e Bit; 
3 km below Garm-e Bit; *Hudar, 26°09’N, 61°27'E; Iran- 
Swiss; 2 km below Iran-Swiss; Kolani marsh, a few km NW 
of Gavater Bay. 


REMARKS.—Apparently the only previously 
published notice of crocodiles in Iran in the zoo- 
logical literature is that of Honegger (1971), 
based on the unpublished work of Bosch, Bul- 
lock, and Kinunen (1970). The Iran Department 
of the Environment in 1971 established the Bahu 
Kalat Protected Region to protect this endan- 
gered species (Firouz 1974:35). This reserve 
covers 394,750 hectares of mountains, foothills, 
and plains, and embraces much of the Sarbaz 
River. The crocodile is a fully protected species 
under Iranian law. However, it faces the double 
threat of hide poaching and habitat destruction. 


ANDERSON: IRANIAN REPTILES 


Y,Emys orbicularis 


SS Agrionemys horsfieldii 
||| Tri onyx euphraticus 


=: Crocodylus palustris 


55 60 65 70 


FiGureE 12. 


Enforcement of protective legislation along the 
Sarbaz River is difficult to impossible at present. 
In 1975 there was no station of the Department 
of the Environment within the Protected Region 
itself, and the office at Chah Bahar had no four- 
wheel-drive vehicles. Without constant patrol of 
the river on horseback, it is doubtful that any 
hide poachers would ever be apprehended. Road 
and bridge construction in the area probably 
poses a threat to the animals, especially if fur- 
ther development results, putting a further de- 
mand on the flow of the river for irrigation and 
domestic use. No adequate study of the croco- 
diles has been carried out following the prelim- 
inary investigations of Bosch, Bullock, and Kin- 
unen. These workers surveyed most of the 
Sarbaz River from the vicinity of Rask to Kolani 
over a two-week period. They actually observed 
a total of 18 individual crocodiles. They con- 
cluded that the total population at that time was 
over 50, but this may have been an optimistic 
figure. The fact that they estimated the maxi- 


Distribution of Emys orbicularis, Agrionemys horsfieldii, Trionyx euphraticus, and Crocodylus palustris. 


mum length of the largest animals seen at about 
two meters may be an indication that hunting 
pressure has eliminated the older mature croc- 
odiles. Their survey indicated that the best croc- 
odile habitat occurs from Firizabad to Djork-e 
Bit and in the Kolani marsh. 


SQUAMATA: AMPHISBAENIA 


Family TROGONOPHIDAE 
Genus Diplometopon Nikolsky 
Diplometopon NikOLsky, 1907:277-280, text-figs. 1-3 (type- 
species: Diplometopon zarudnyi Nikolsky, 1907, by 
monotypy). 


DEFINITION.—Without limbs; pectoral annuli 
not differentiated from other annuli; frontal di- 
vided or partially divided by a longitudinal sul- 
cus, not in contact medially; eyes conspicuous; 
no lateral line, nor posterior fold; deep ventral 
median sulcus extending almost entire distance 
from head to anal region; tail acuminate; preanal 
pores present. A single species. 


524 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41, No. 22 


Mauremys caspica 


4 caspica 


SN. rivulata 


FiGur_E 13. 


REMARKS.—See Gans (1960) for a review of 
the trogonophid amphisbaenians. 


Diplometopon zarudnyi Nikolsky 
(Figures 11, 14) 


Diplometopon zarudnyi NIKOLSKY, 1907:277-280, text-figs. 1- 
3 (type-locality: Nasrie, Arabistan, Iran; holotype: Zoolog- 
ical Institute, Leningrad no. 10341). —WERNER 1936:200. 
—GAaNns 1960: 133-204, pl. 45, figs. 1, 3, 5-8, 12-14, 18, 20, 
22-24, 29-30. —ANDERSON 1963:456. —GANS 1967:67; 
1968:345-362. —ANDERSON 1968:333. —Tuck 1971:60. — 
ANDERSON 1974:30, 44. 


DiaGnosis.—As for the genus. 

CoLor PATTERN.—Dorsum light gray (pink- 
ish gray in life), heavily flecked with black over 
entire length of back; head and venter dirty 
white (pink in life). 

SIZE.—Snout-vent length 187 mm, tail 15 mm. 

HABITAT.—I collected a single specimen on 
the Ahvaz plain in Khuzestan in an active dune 
area with sparse vegetation. It was a few cen- 
timeters below the surface. I could not deter- 


Distribution of Mauremys caspica. 


mine whether it had been in a burrow or was 
moving through loose sand. 
DISTRIBUTION.—The distribution of this 
species centers about the head of the Persian 
Gulf, in low, sandy deserts of southwestern 
Iran, southern Iraq, Kuwait, the northern half 
of Saudi Arabia, and the Trucial States. 
ZOOGEOGRAPHIC ELEMENT.—Saharo-Sindian. 


IRANIAN LOCALITIES.—Ostan 6 (Khuzestan—Lorestan): 
*Ahvaz (USNM; CG 60; RT 71); *31°16’N, 49°11’E, dunes on 
road between Ahvaz and Haft Gel (CAS; SA 63); Nasrie [very 
close to Ahvaz, on Rud-e Karun] (AN 07; type-locality; CG 
60). 


REMARKS.—Diplometopon zarudnyi has its 
nearest living relatives in the species of Aga- 
modon and Pachycalamus among the Trogo- 
nophidae. The morphology, adaptations, and re- 
lationships of this form are discussed in detail 
by Gans (1960). This species should be looked 
for in suitable dune habitats in Fars Province 


ANDERSON: IRANIAN REPTILES 


525 


MEDITERRANEAN 


See 


W/ 


ee 
SEA 
f -* . 
ye ae 
(B fe f ee: -- 


Testudo graeca 
UY, ibera 
S zarudnyi 


\\|| terrestris 


45 


4 
Fast 


40 


FiGurReE 14. 


along the coast of the Persian Gulf. It is part of 
the gulf coast fauna that extends south to Bu- 
shire, and perhaps as far south as 27°50’N. Con- 
tinued survival of this fauna will depend upon 
action to preserve dune habitat in the face of 
pressure for dune stabilization. 


MATERIAL EXAMINED 


I list here only Iranian specimens. In addition, 
however, I have examined most of the reptiles 
from elsewhere in Southwest Asia in the Amer- 
ican Museum of Natural History, California 
Academy of Sciences, Field Museum of Natural 
History, United States National Museum, Uni- 
versity of Michigan Museum of Zoology, and 
University of California at Berkeley Museum of 
Vertebrate Zoology. I have listed specimens 
only by political provinces (ostans); see text for 
specific localities from which I have seen 
material. 

The collectors of the bulk of the material ex- 


Distnbution of Testudo graeca and Diplometopon zarudnyi. 


amined are as follows: Steven C. Anderson, 
1968: CAS 86514-86639; William S. and Janice 
K. Street Expedition to Iran 1962-63: FMNH 
141619-141631, CAS 102481-102483; Robert G. 
Tuck, Jr., John W. Neal, Jr., G. L. Ranck, and 
L. H. Hermann, 1962-65: USNM 153752- 
158945; Street Expedition to Iran 1968: FMNH 
170989—170993; Steven C. Anderson, Robert G. 
Tuck, Jr., Ruben McCullers, Reza Khazai’e, 
1975: MMTT 978-1530, CAS 141166—141236. 


Eretmochelys imbricata bissa: PERSIAN GULF: MMTT 978— 
979. 

Emys orbicularis: EAst AZARBAWAN: MMTT 1278; MAZAN- 
DERAN: AMNH 88424; MMTT 401-402; FMNH 141626. 
Mauremys caspica caspica: GiLAN: FMNH 141619, 141627; 
USNM 154504-154505; East AZARBAIJAN: MMTT 47, 2 live 
spec.; FMNH 141625, 141629; CAS 102483; KorDESTAN-— 
KERMANSHAH: FMNH_ 170989; USNM_ 153752, 158529- 
158534; MMTT live spec.; KHUZESTAN-LORESTAN: CAS 
86629, 86634-86639, 141118; USNM 158945; FMNH 
130819, 73492, 73495; FArs: FMNH 21035-21042, 141624; 
CAS 102481-102482; MMTT 208; MAZANDERAN: MMTT 

421-422, 492, 1232; AMNH 88662; CAS 141166. 


526 


Agrionemys horsfieldii: MAZANDERAN: MMTT S33. 

Testudo graeca ibera: TEHRAN (CENTRAL): CAS 143300: GI- 
LAN: FMNH_ 170990-170991; East AZARBAUJJAN: MMTT 
493, 1446, live spec.; West AZARBAIJAN: FMNH 141630- 
141631; MMTT 555; KorDESTAN-KERMANSHAH: FMNH 
130820, 170992-170993; MMTT 1530, 3 live spec.; KHU- 
ZESTAN—LORESTAN: FMNH 73483-73484, 73487-73490; 
EsFAHAN: FMNH 141620; FArs: FMNH 21027-21029. 

Testudo graeca zarudnyi: BALUCHISTAN-SISTAN: MMTT, 3 
live spec. 

Diplometopon zarudnyi: KHOZESTAN—LORESTAN: CAS 86514; 
USNM 121594. 


Maps 


The distribution in Southwest Asia is given 
for each species; only the Iranian localities are 
shown. Each symbol represents one or more 
documented locality, as listed in the text. Stars 
represent type-localities. Solid symbols repre- 
sent localities from which I have identified and 
examined specimens. Open symbols indicate lo- 
calities based on literature or museum cata- 
logues only. 


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SMITH, HOBART M., AND EDWARD HARRISON TAYLOR. 
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199: 1-253. 

SmITH, MALCOLM A. 1931. The fauna of British India. Rep- 
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Francis, London. xxviii + 185 pp., 2 pls. 

SOBOLEVSKY, N.I. 1929. The herpetofauna of the Talysh and 
of the Lenkoran lowland. Mem. Soc. Amis Sci. Nat. An- 
thropol. Ethnog. Sect. Zool. 5:1—-141. [In Russian; English 
summary.] 


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STRAUCH, ALEXANDER. 1862. Essai d’une erpetologie de 
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TERENTJEV, PAUL V., AND S. A. CHERNOV. 1949. Oprede- 
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THUNBERG, CARL PEHR. 1787. [Not seen; cited in Wermuth 
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Tuck, Rosert G., Jr. 1971. Amphibians and reptiles from 
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WAGLER, JOHANN GEORG. 1830. Nattrliches System der 
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WALCZAK, PETER S. 1971. Green sea turtle nests and turtle 
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C953 


INDEX TO VOLUME XLI 


FOURTH SERIES 


New names and principal references in boldface type. 


Generic and specific names occurring in No. | are indexed on pages 120-123 of that number; they are not 
included in the following index. 


Ablepharus bivittatus 510 
Abyssicola 310-311 
macrochir 310-311 
Acacia 521 
Acanthosphex 404 
leurynnis 404-405 
Achnanthes 
brevipes 341 
coarctata 341 
grimmei 341 
hungarica 341 
inflata 341 
kryophila 341 
lapponica 341 
linearis 341 
marginulata 341 
minutissima 341 
pyrenaica 341 
trinodis 341 
Acila castrensis 364 
Adula californica 364 
Adventor 404 
elongatus 404—405 
Agamodon 524 
Agavaceae 371 
Agave 371-386 
aurea 374-376, 380-382, 384-385 
avellanidens 374-378, 384, 386 
capensis 375-376, 382-385 
cerulata 374, 376-377, 379, 384-385 
cerulata cerulata 375-378, 384 
cerulata nelsonii 375-378, 386 
cerulata subcerulata 375-378, 384 
deserti 372, 374, 376, 379, 384-385 
deserti deserti 375, 376-378, 384 
deserti pringlei 375, 376-378 
deserti simplex 375, 376-378, 384 
ellemeetiana 384 
fortiflora 371, 380-382, 385 
gigantensis 374-376, 377-379, 384-386 
margaritae 375-379, 384-385 
mcekelveyana 375-376, 379-381, 384 
moranii 374-376, 379-381, 384, 386 
promontorii 375-376, 382, 383-386 
schottii 386 
sebastiana 375-376, 382, 383-384, 386 
shawii 382, 384, 386 
shawii goldmaniana 373-376, 382, 383-384, 386 
shawii shawii 375-376, 383-384 


sobria 374, 384 

sobria frailensis 373, 375-376, 379-381, 384-386 

sobria roseana 372-373, 375-376, 379-381, 385- 

386 

sobria sobria 373, 375-376, 379-381, 384 

subsimplex 373-376, 379-381, 384-385 

victoriae-reginae 386 

vizcainoensis 373, 375-376, 379-382, 384-386 
Agrionemys 501, 513 

horsfieldii 501, 505, 513-514, 518, 523, 526 


Alepisaurus 243 


Amblycheila schwarzi 180 
Amphipoda 229 
Amphisbaenia 523 
Amphissa reticulata 364, 368-369 
Amphora coffeaeformis 341 
Amyda euphratica 519 
Anadara trilineata 301 
Aniculerosa 404—405 

taprobanensis 401, 403-405, 416, 419 
Anomoeoneis 

styriaca 341 

zellensis 341 
Anoplogasteridae 231 
Apistes 461 

monodactylus 461 

Russellii 461 
Apistus 

dermacanthus 405 

minous 461 

monodactylus 461 

niger 479 
Aploactidae 402 
Aploactinae 402 
Aploactinidae 401-405 
Aploactis 404 

aspera 401, 404-405, 419 

aspersa 419 

lichen 405, 411 

milesii 405, 420 

pusillus 462 

trachycephalus 464 
Aploactisoma 404-405 

milesii 405 

milesii horrenda 405 

schomburgki 404—405 
Aploactoides 404 

philippinus 404-405 
Aploactus sieboldi 405 


[529] 


530 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41 


Apoidea 281 
Artemisia 517 
Arthropoda 364 
Asilidae 183, 186, 196, 207 
Astrodapsis 298 

n.sp. 298 

perrini 297 
Ataxolepis 230 


Balanus 

sp. 364, 386 

(Tamiosoma) gregarius 300 
Barbourisiidae 231 
Bassogigas 390-391, 398 

gillii 390 

grandis 390 
Bathyaploactidae 402 
Bathyaploactis 404 

curtisensis 404-405 

curtisensis ornatissimus 405 
Beryciformes 227, 230-231 
Bittium sp. 364, 368-369 
Bivalvia 364 
Bohartia 188 
Bryozoa 364 


Caenopangonia 427-428, 446, 449-451 
asper 427, 446-448, 450, 452 
brevirostris 427, 441, 446-448, 450-452 
hirtipalpis 427, 446-448, 450-451 

Calanoida 229 

Callianassa sp. 364 

Caloneis 
clevei 341 
ladogense var. densestriata 341 
obtusa 341 

Calyptraea filosa 364 

Campaniflorae 371-372, 374-376, 382, 385, 387 

Cancellaria 
palmeri 298 
sp. 364 

Cancer sp. 363-364 

Caretta 504, 506 
bissa 507 
caretta 506 
caretta gigas 501, 505-506 
gigas 506 
nasuta 506 

Cetomimidae 231 

Cetomimiformes 215, 228, 230 

Cetomimoidei 231 

Cetonurus 235 

Cetunculi 230 

Chelonia 504, 506 
mydas 505, 506 
mydas japonica 501, 505, 506—507 
olivacea 507 

Chelontidae 506 


Chitra 519 
Choridactylinae 453, 475-477, 495 
Choridactylodes 475, 479 
natalensis 479, 482 
Choridactylus 453, 475-479, 481-482, 494, 496 
multibarbis 479 
multibarbus 477-478, 479-483, 494 
natalensis 477-481, 482-483, 494 
Chorismodactylus 475, 479 
multibarbis 479-480 
multibarbus 479 
Chorismopelor 475-476, 483, 492 
Joubini 483, 492 
Chrysothamnus paniculatus 282-283 
Cicindela 169-170, 176, 178-179, 181 
beneshi 181 
carthagena carthagena 176 
celeripes group 179 
(Cicindela) 178, 181 
circumpicta group 179 
(Cylindera) 179 
debilis 177-180 
debilis segnis 177 
(Ellipsoptera) 179 
fulgida 179-181 
(Habroscelimorpha) 179 
haemorrhagica 176-179 
haemorrhagica woodgatei 176, 178 
hirticollis 179 
hirticollis ponderosa 179 
horni 178-180 
horni horni 176-177, 179 
lemniscata 177-180 
lemniscata group 179 
lemniscata lemniscata 177, 179 
lepida 179 
marutha 176-179 
marutha marutha 176, 179 
marutha rubicunda 176 
nevadica 175, 178-180 
nevadica citata 169, 173, 175-176 
nevadica group 179 
nevadica knausi 176 
nevadica nevadica 175-176 
nevadica tubensis 176 
nigrocoerulea 178-180 
nigrocoerulea nigrocoerulea 176, 179 
obsoleta 178-180 
obsoleta group 179 
obsoleta santaclarae 177-179 
obsoleta vulturina 180 
ocellata 178-180 
ocellata ocellata 177-178 
ocellata rectilatera 177, 180 
parowana 181 
pimeriana 177-178, 180-181 
praetextata 178-180 
praetextata erronea 176, 179-180 


INDEX 


praetextata fulgoris 176, 180 
praetextata praetextata 176, 180 
pulchra 174-175, 177-181 


pulchra dorothea 169, 172-175, 177-178, 180-181 


pulchra group 178, 180-181 

pulchra pulchra 174-175, 181 
punctulata 178-180 

punctulata chihuahuae 176, 178, 180 
punctulata group 178 

rockefelleri 181 

rufiventris group 178 
sedecimpunctata 178-180 


sedecimpunctata sedecimpunctata 177-178 


spaldingi 172 
sperata 179 
tenuisignata 176, 178-180 
viridisticta 178 
viridisticta arizonensis 178-180 
willistoni 172, 178-180 
willistoni amadeensis 172 
willistoni echo 172 
willistoni estancia 172 
willistoni funaroi 172 
willistoni group 178 
willistoni hirtifrons 172 
willistoni praedicta 172 
willistoni pseudosenilis 172 
willistoni sulfontis 169, 170-173, 177-178 
willistoni willistoni 172 
Cicindelidae 169, 179 
Cicindelidia 178 
Cistudo europaea 510 
Clemmys 511 
caspica 511 
caspica caspica 511 
Clinocardium 
meekianum 363-364 
nuttallii 300 
sp. 364 
Cocconeis disculus 341 
Coccotropus obbesi 404-405, 419-420 
Cockerellia 281-282 
Cocotropus 401, 404, 407-408, 419 
abbesi 419 
altipinnis 404-405, 407 
dermacanthus 405, 407 
de Zwaani 405 
dezwaani 416 
echinatus 405, 407 
hongkongiensis 401, 405, 416, 420 
kagoshimensis 405, 417 
masudai 404—405, 407, 419 
monacanthus 405, 407 
obessi 401, 416 
pottii 404—405, 416 
roseus 405, 407 
steinitzi 405, 407 
(Tetraroge) roseus 405 


531 


Coelorhynchus 
canus 317 
chilensis 332 
(Coelorhynchus) canus 317 
(Coelorhynchus) patagoniae 322 
(Coelorhynchus) scaphopsis 313 
fasciatus 322 
innotabilis 329 
(Oxymacrurus) chilensis 332 
(Paramacrurus) fasciatus 322 
(Paramacrurus) innotabilis 329 
patagoniae 319, 322 
scaphopsis 313 
Coelorinchus 307, 309, 310-313, 315-319, 321-322, 
325, 329, 332, 334-335 
aconcagua 307, 311, 313, 315-316, 319-322, 328, 
Bs s35 
aspercephalus 311, 335 
asteroides 316 
australis 335 
canus 307, 311-313, 315-316, 317-319, 335 
caribbaeus 312, 315, 319, 335 
chilensis 307, 311-313, 315-316, 328, 331, 332-335 
cingulatus 335 
coelorhincus 311-312, 316, 335 
coelorhincus carminatus 312, 315, 331 
(Coelorinchus) 311 
fasciatus 307, 311-313, 315-316, 321, 322-329, 
33)Il, 3333)5 3335) 
hubbsi 321 
innotabilis 307, 313, 315, 328, 329-333, 335 
labiatus 312 
La-Ville 310 
mirus 328, 335 
mortoni 335 
occa 311-312, 322, 333-334 
oliverianus 321, 335 
(Oxygadus) 311, 333-335 
(Oxymacrurus) 311, 333-335 
(Oxymacrurus) japonicus 334 
(Oxymacrurus) tokiensis 334 
(Paramacrurus) 307, 311, 335 
patagoniae 307, 316, 321, 328 
(Quincuncia) 311 
scaphopsis 307, 311-312, 313-317, 319, 335 
smithi 330, 333 
tokiensis 325 
velifer 312 
ventrilux 335 
Coleoptera 169, 188, 346, 355 
Compsomyax subdiaphana 364 
Congiopodus leucometopon 408 
Copepoda 229 
Coryphaenoides 334 
acrolepis 234 
armatus 233 


filifer 233 


532 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41 
leptolepis 233 Denticula 
rupestris 234 tenuis 341 
Corythobatus 453-454, 461 vanheurcki 342 
echinatus 404—405 Dermochelyidae 507 
monodactylus 461 Dermochelys 503, 507, 509 
trachicephalus 464 coriacea 501, 504-505, 507, 509 
woora 461 Deserticolae 371, 374-376, 385-386 
Crepidula Diatomella balfouriana 341 
princeps 363 Diatomineura hirtipalpis 428, 446, 452 
aff. princeps 364, 368-369 Diclisa misera 354 
Crocodilia 520 Dioctriini 188 
Crocodilus palustris 520 Diplometopon 523-524 
Crocodylidae 520 zardnyi 501-502, 522-523, 524-526 
Crocodylus 520 Diploneis 
niloticus 520 domblittensis 342 
palustris 501, 520-523 domblittensis var. subconstricta 342 
palustris brevirostris 522 elliptica 342 
palustris palustris 520-522 marginestriata 342 
Cryptomya californica 364, 368-369 mauleri 342 
Cyclotella ovalis 342 
distinguenda 341 pulcherrima 342 
elgeri 341 Diptera 183, 186, 345, 427, 451 
iris 341 
stelligera 341 Echinoida 364 
Cylindera 179 Echinoidea 297 
Cymbella Echinomacrurus 233, 235-236, 250 
alpina 341 mollis 234 
aspera 341 Ellipsoptera 179 
bernensis 341 Emys 501, 509 
botellus 341 caspia 511 
cistula var. arctica 341 europaea 509 
gracilis 341 fulginosa 510 
hebridica 341 lutaria 510 
lapponica 341 orbicularis 501, 505, 509-510, 523, 525 
norvegica 341 Epithemia 
ruttneri 341 sorex var. lapponica 342 
sumatrensis 341 turgida 342 
turgidula 341 Epitonium 
Cynomacrurus 235-236, 250 sp. 364 
piriei 233-234, 236, 238, 250 tinctum 364 
Cyphocaris johnsoni 229 Eretmochelys 504, 507 
imbricata 504, 507 
Dactylagnus 127, 129-130, 150 imbricata bissa 501, 504-505, 507-508, 525 
Dactyloscopidae 125, 130 Erisphex 404, 416-417 
Dactyloscopus 126, 129-130, 155 achrurus 405 
cinctus 125, 154-155, 157, 160 kagoshimensis 416-417 
lunaticus 137 obbesi 419 
Dasylirion 386 Philippinus 405 
Dasypogoniae 186—187 potti 404-405, 416 
Decterias 453-455, 462 pottii 404-405 
pusillus 462 Erosa australiensis 403, 405, 411 
Dendraster 297-298, 303, 305, 363 Esenbeckia 345-347, 350, 353 
arnoldi 303 abata 345, 349 
elsmerensis 303 basilaris 351 
excentricus 303 breedlovei 351 
gibbsii 301, 303 caustica 354 
perrini 297-298 curtipalpis 352 
sullivani 297, 303-305 delta 345, 347, 353-354 


vizcainoensis 303 deltachi 347, 353 


INDEX 33/3) 


divergens 350 semiflava melanista 351 
downsi 352 semiflava var. melanista 348 
(Esenbeckia) 350, 353 semiflava melanopa 351, 353 
(Esenbeckia) divergens 347 semiflava subspp. 351 
(Esenbeckia) illota 350 seminuda 353-354 
(Esenbeckia) minor 347 tepicana 345, 351, 354 
(Esenbeckia) nigronotata 350 tepicana abata 351, 354 
(Esenbeckia) wiedemanni 347-348 translucens 350 

fascipennis 347 triangularis 353-354 
fidenodes 352, 354 weemsi 351 

filipalpis 347 wiedemanni 348, 350, 354 
flavohirta 352 Eucalanidae 229 

fuscipes 350 Eucalanus elongatus 229 

gertschi 351, 354 Eunotia 

hoguei 352-353 arcus 342 

illota 350 bigibba 342 

incerta 353-354 clevei 342 

incisuralis 345, 347, 353-354 denticulata 342 

incisuralis tinkhami 347 lapponica 342 

keelifera 352 serpentina 342 

leechi 351 triodon 342 

mejiai 347, 352, 354 Eutaeniophoridae 231 

melanopa 345, 348 Evermannella ahlstromi 243 
mexicana 352 

micheneri 347, 350, 353 Fidena 355, 448 

nigronotata 353-354 flavipennis 346, 355-356 
nitens 352 flavipennis fisheri 345, 355-356 
painteri 351 isthmiae 356 

pavida 351, 353 Fragilaria 

planiventris 349, 352, 354 capucina 342 

var. planiventris 352 crotonensis 342 

planiventris saussurei 354 javanica 342 

planiventris var. saussurei 349, 352 

prasiniventris 350 Gadiformes 233 

(Ricardoa) 345-346, 348-350 Gammaridea 229 

(Ricardoa) caustica 349 Garichthys fasciatus 322 
(Ricardoa) curtipalpis 350 Gastropoda 364 

(Ricardoa) deltachi 345, 346-347 Gecko 

(Ricardoa) downsi 350 listeri 258 

(Ricardoa) flavohirta 350 moestus 258 

(Ricardoa) incerta 350 pumilus 262 

(Ricardoa) incisuralis tinkhami 347 Gehyra 263 

(Ricardoa) incisuralis var. tinkhami 350 Gekko 258-259, 262—264 
(Ricardoa) leechi 345, 346 oorti 261 

(Ricardoa) nigronotata 350 pumilus 261 

(Ricardoa) painteri 350 Gekkonidae 253 

(Ricardoa) pavida 350 Gekkoninae 263 

(Ricardoa) planiventris 349-350 Gibberichthyidae 215-216, 227, 230-231 
(Ricardoa) schusteri 349 Gibberichthys 215, 226-228, 230 
(Ricardoa) scionodes 350 latifrons 215, 219, 222-225, 227-229 
(Ricardoa) semiflava melanista 348 pumilus 215-230 

(Ricardoa) semiflava melanopa 348 Gillellus 125-126, 127-132, 136-137, 141-143, 145, 
(Ricardoa) seminuda 350 149-150, 154, 158 
(Ricardoa) tepicana 349 arenicola 125, 127, 131-132, 134-137, 140-143, 
(Ricardoa) weemsi 350 145-146, 149, 158 

saussurei 349-350 australis 125-126, 150, 154 
schusteri 351, 353 chathamensis 125, 127, 131, 136-137, 138-143, 
scionodes 352, 354 145-146, 149, 158 


semiflava 348, 351, 353 greyae 150 


534 


ornatus 125, 127, 
144-146, 149, 158 
searcheri 125-126, 128, 131-132, 134, 136-137, 
141-143, 145, 146-150, 158-159 
searcheri group 150 
semicinctus 125, 127-128, 131, 132-137, 139, 141- 
143, 145-146, 149, 158-159 
Gomphocymbella ancyli 342 
Gomphonema 
cantalicum 342 
eriensee 342 
quadripunctatum 342 
subtile 342 
transilvanicum 342 
ventricosum 342 
Gopherus 513 
Gymnodactylus caudeloti 258 
Gymnothorax 163 
breedeni 161, 162—165 
flavimarginatus 164 
monostigma 164 
sp. 162 


131-132, 135-137, 141-143, 


Habroscelimorpha 179 
Harpacticoida 229 
Hemidactylus meijeri 258 
Hemiphyllodactylus 263 
Heteristius 125-126, 128-132, 136-137, 141-145, 149, 
154-155, 158 
cinctus 127-128, 131-132, 134, 136-137, 141-143, 
145, 148-149, 152, 154, 155-158 
Jalisconis 125, 154-155, 157 
Hexaperdita 281, 283-284 
Hoplistomerus 186-187 
engeli 187 
nobilis 186-187 
zelimina 187 
Hymenocephalus 236 
Hymenoptera 188, 281 
Hyperiidea 229 
Hyperopsidae 229 
Hyperiopsis sp. 229 


Idiacanthus 235 
Inimicinae 475—477 
Inimicus 453, 475-478, 481, 483-484, 489, 492, 494— 
496 
auranticus 493, 495 
barbatus 488-490 
bifilis 475, 484, 490 
brachyrhynchus 475, 478-479, 481, 484, 491-492, 
494 
caledonicus 475, 477—478, 481, 484, 488-489, 494 
cirrhosus 475, 487, 490-491 
cuvieri 475, 477-478, 481, 484-485, 486-488, 490, 
494, 496 
didactylum 490 
didactylus 475, 477-478, 481, 484, 488, 489-491, 
494-497 


PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41 


filamentosus 475-478, 481, 484-486, 494, 496 
japonicus 475, 478-479, 481, 484, 492, 493-495 
Japonicus var. auranticus 495 
joubini 475, 478-479, 481, 484, 492-494 
(Simopias) 483 
sinense 487 
sinensis 475, 477-478, 481, 484-485, 487-488, 494 
Insecta 183, 345 
Insopiscis 404 
Tracundus 496 
signifer 496 
TIsocladus sp. 154 


Kanekonia 404 
aniara 404—405 
florida 404—405 
queenslandica 405 
Karumba 404 
ornatissima 404—405 
Kasidoridae 215, 228 
Kasidoroidae 215, 228 
Kasidoron 216, 228, 230 
edom 215-217, 219, 223-226, 228, 230 
latifrons 215-216, 219, 223, 226, 228 
Kleiwegia 404 
dezwaani 404 


Lacerta 510 
saxicola 510 
Lampridiformes 231 
Laphriinae 183, 186-188 
Laphriini 186 
Laphystia 183, 188, 190, 196 
carnea 187 
jJamesi 189-191 
litoralis 190-191 
sabulicola 188 
spp. 190-191 
Laphystiini 183, 186-189, 194, 196 
Lepidochelys 504, 507 
olivacea 501, 505, 507 
Lepidodactylus 253-265 
divergens 258 
gardeneri 254-256, 258, 264 
guppyi 254-255, 257-258, 263-264 
guppyi-pulcher group 263 
listeri 254, 256, 258, 264 
lugubris 254-255, 257-258, 264 
lugubris-woodfordi complex 253, 255, 264 
magnus 253-254, 256-257, 258-259, 263-264 
manni 254, 256, 259, 264 
mutahi 253-254, 256, 259-260 
novaeguineae 253-255, 260-261, 263-264 
oorti 254, 256, 261, 264 
orientalis 253-254, 256, 261-264 
' pulcher 254-255, 257, 260-262, 264 
pumilus 254, 256-257, 262-265 
pumilus-oorti group 263 
shebae 254, 256, 262-264 


INDEX 


species 263 

woodfordi 261, 263—264 
Lepidoptera 454 
Leurochilus 129, 132 
Lithothamnion 420 
Luperosaurus 263 
Lysianassidae 229 
Lysodermus 453-454 

satsumae 454, 461 


Macoma 
inquinata 364, 368-369 
nasuta 363-364, 368-369 
sp. 364 
Macrouridae 233, 236, 311-312 
Macrourinae 311 
Macrouroides 250 
inflaticeps 234, 250 
Macrurus 
canus 317 
(Coelorhynchus) canus 317 
(Coelorhynchus) fasciatus 322 
(Coelorhynchus) scaphopsis 313 
fasciatus 322 
fragilis 242 
trichiurus 242 
Malacocephalus 240, 242-243 
sp. 243 
Martinia 187 
Mauremys 501, 510-511 
caspica 505, 510-511, 516, 524 
caspica caspica 501, 505, 511-512, 524-525 
caspica rivulata 524 
Megalomycter 230 
Megalomycteridae 215, 228, 231 
Melamphaeidae 230-231 
Melosira 
arenaria 342 
ruttneri 342 
Membracidichthys 404—405 
(Adventor) elongatus 405 
obbesi 404, 419 
Merriamaster 295-298, 301 
arnoldi 301, 303 
israelskyi 303 
kewi 303 
pacificus 303 
perrini 297-298, 301, 303 
ef. perrini 303 
weaveri 297-298, 301-303 
Mesembryanthemum 
crystallinum 212 
Mesobius 233-234, 235-236, 238, 240, 242-243, 245, 
250 
antipodum 233, 236, 242-244, 245-246 
berryi 233, 235, 236-246 
Metridiidae 229 
Microstomus 
pacificus 312 


535 


Minaus pusillus 462 
Minoinae 453-454, 476, 495 
Minois 454 
Minous 453, 454-456, 458 
adamsi 460-461 
adamsi var. inermis 458 
adamsii 461 
Blochi 461 
coccineus 456-458, 464, 465-467 
dempsterae 453, 456—458, 468, 469-470 
echigonius 461 
inermis 454-458, 463-465 
longimanus 463-464 
longipinnis 463-464 
monodactylus 454-459, 460-462, 466 
oxycephalus 461 
oxyrhynchus 461 
pictus 456, 458, 464, 467-469 
pusillus 454-458, 462-463 
quincarinatus 455-457, 458-460, 463 
satsmae 461 
satsumae 461 
superciliosus 464, 466 
trachycephalus 455-458, 463, 464-467 
versicolor 455460 
woora 461 
Mirapinnatoidei 215, 228 
Mirapinnidae 231 
Mirapinniformes 215, 230 
Mirounga angustirostris 267-279 
Mitrella gouldii 363-364 
Modiolus sp. 364 
Muraenidae 161 
Mycteromyia 427-428, 433, 441, 443, 448, 450-452 
asper 428, 448 
bejaranoi 427, 446, 452 
brevirostris 428, 446 
cinerascens 435, 439, 445 
conica 427-428, 430, 441-443, 449, 452 
edwardsi 446 
eriodes 434 
etcheverryae 427, 440-441, 443, 449, 452 
flaviventris 445 
fusca 428, 441 
martinezi 445 
murina 438 
Philippii 428, 430, 441 
robusta 427, 448, 452 
Mycteromyiini 427—428, 429-430, 443, 448, 450-452 
Mytilus 
coalingensis 364 
condoni 363-364, 368-369 
highoohiae 363 
sp. 364 
Myxodagnus 127, 129-130, 142, 150 


Nassarius 357 
(Caesia) grammatus 300 
mendicus 364 


536 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41 


moranianus 363-364, 368-369 
sp. 364 
Natrix tessellata 510 
Navicula 
accommoda 342 
grimmei 342 
Jakovljevici 342 
Kincaidii 339 
ludloviana 343 
perpusilla 343 
placentula 343 
pseudobryophila 343 
tenuicephala 343 
Neidium 
distinctepunctatum 343 
hitchcocki 343 
meisteri 343 
Neoaploactis 404 
tridorsalis 404—405 
Neobythites 398 
grandis 390 
Neophron percnopterus 512 
Neptunea tabulata 364 
Nerium 522 
Nezumia 234, 240, 246, 249, 334 
aequalis 249 
africana 249 
atlantica 249 
bubonis 249 
condylura 249 
convergens 243 
holocentrus 249 
liolepis 239, 249, 317 
parini 233, 236, 243-244, 246-250 
pudens 249 
sp. 246 
stelgidolepis 249 
Nolina 386 
Nucella | 
canaliculata 364, 368-369 
emarginata 364 
lamellosa 363-364 


Odontomacrurus 235-236, 242-243, 250 
murrayi 233-234, 236, 238, 240, 242, 250 

Olivella 
biplicata 363-364 

Ophidiidae 398 

Ophiodermella 357, 363, 369 
graciosana 357, 364, 368-369 
graciosana var. mercedensis 369 
incisa 357, 368-370 
mercedensis 357, 363-364, 368-370 
sp. 364 

Orthoneuromyia 190 
modesta 190 

Oxygadus 311, 333-335 

Oxymacrurus 311, 333-335 


Pachycalamus 524 
Pandanus 259 
Pangonia 
conica 427-428, 441 
obscuripennis 427, 433, 452 
Pangoniinae 345, 427, 450-452 
Pangoniini 449 
Parabassogigas 390, 398 
grandis 389-398 
Paracetonurus 235, 243 
Paramacrurus 307, 311, 335 
Paraminous 453-455, 463 
inermis 463 
quincarinatus 454, 458, 463 
Paraploactis 401-404, 405-412, 414-416, 420 
hongkongiensis 405, 419-420 
intonsa 401, 404-413, 414-416, 425 
kagoshimensis 401, 404-405, 416-419 
kagoshimensis complex 401, 406-413, 416-417, 
426 
obbesi 405, 418, 419-420 
pulvinus 401, 406-412, 413-415, 423 
species 419-420 
taprobanensis 406, 418-419 
trachyderma 401, 404-410, 411-414, 424 
Patinopecten 
healeyi 300 
(Lituyapecten) 297, 301 
(Lituyapecten) falorensis 297, 300-301 
(Lituyapecten) purisimaensis 297, 300-301 
(Lituyapecten) turneri 297, 300 
lohri 301 
Pelecorhynchidae 428, 451 
Pelochelys 519 
Pelor 475, 477, 483 
auranticum 493, 495 
barbatus 488-489 
brachyrhynchus 491 
caledenicum 484, 488-489 
cuvieri 486—487, 491 
didactylum 484, 488-489 
didactylus 490 
filamentosum 483-484, 486 
filamentosus 484 
Japonicum 483, 493 
maculatum 489-490 
obscurum 483, 489-490 
sinense 486—487, 489 
‘tigrinum 493 
Pelorinae 475, 477 


Perasis 183, 188, 192, 194, 196 


argentifacies 189, 194-196 
sareptana 192, 194 
transvaalensis 192 


Perdita 281, 286, 290 


albipennis 282 
apicalis 281, 290, 292-293 
aridella 289 


INDEX 


autumnalis 281—282 
bifasciata 281, 290, 293-294 
bishoppi 283 
boltoniae 283 
cazieri 289 
cinctiventris 281, 283 
(Cockerellia) 281-282 
cushmani 292 
dammersi 292 
diminutiva 281, 291 
ensenadensis 286 
esmeraldensis 281, 286—287 
exilis 291 
falcata 292 
foveata 283 
gracilior 281, 290, 291-292 
graenicheri 283 
heterothecae 284-285 
(Hexaperdita) 281, 283-284 
infuscata 281, 283-284 
inyoensis 281, 287 
koebelei 292 
lingualis 282 
lucidella 294 
luculenta 281 
medialis 281, 286, 287-289 
minima 285 
obliqua 292 
pachygnatha 281, 282—283 
pauliana 281, 289-290 
(Perdita) 281, 286 
(Perditella) 281, 285 
perlucens 281, 294 
polygonellae 287 
pusillissima 281, 285-286 
sejuncta 286 
semicrocea 291 
swenki 287, 289 
willcoxiana 281, 289-290 
zavortinki 281, 284—286 
zebrata 293 
Perditella 281, 285 
Periophthalmus sp. 521 
Peripia 
mysorensis 258 
ornata 258 
Peristromindus 404 
dolosus 404—405 
Perochirus 263 
Peronia heribaudi 343 
Peropus neglectus 258 
Phalacromacrurus pantherinus 242-243 


Pinnularia 
balfouriana 343 
gracillima 343 
polyonca 343 
pulchra 343 


357/ 


semicruciata 343 
transversa 343 

Pisces 125, 161, 233 

Platydactylus lugubris 258 

Platygillellus 127, 129-130, 132, 150, 155 

Pleuromamma abdominalis 229 

Polinices lewisii 363-364 

Poromitra 230 

Prionace glauca 389 

Promycteromyia 427—428, 430, 439, 448, 450-452 
cinerascens 427, 430, 433-434, 435-436, 438, 449_- 

450, 452 

derocerca 427, 430, 432, 433-434, 438, 450, 452 
eriodes 427, 430, 433, 434-435, 449, 452 
galbina 427, 430, 431-433, 435, 438, 450, 452 
murina 427, 430, 433, 437, 438-439, 450, 452 
pechumani 427, 430, 436-438, 449, 452 
penai 427, 430, 437, 438-439, 449, 452 
philippti 427, 429, 430-431, 433, 449, 452 
xantha 427, 430, 439-441, 449, 452 

Prosopis 521 

Prosopodasys asperrimus 405 

Protothaca tenerrima 364 

Pseudogekko 263 
shebae 262 

Psilocurus 183, 187-188, 190, 192 
nudiusculus 190 
sp. 189, 192-193 

Ptarmus gallus 408 

Pteroinae 404 

Ptychozoon 263 


Quincuncia 311 


Rana 
camerani 510 
cyanophlyctis 521 
ridibunda 512 
Ricardoa 350 
Rondeletia 230 
bicolor 230 
Rondeletiidae 231 
Rosauridae 231 


Sargassum 230 
Saucropogon 192, 194 
Saxidomus sp. 364 
Scincidae 264 
Scione 354 
aurulans 345-346, 354-355 
lurida 355 
Scionini 427-428, 446, 451 
Scorpaena 
biaculeata 461 
didactyla 483, 489-490 
digitata 489-490 
monodactyla 454, 460-461 
Scorpaenidae 402, 453, 476-477 


538 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Ser. 4, Vol. 41 


Scorpaeniformes 401 

Scorpaeninae 495 

Scorpaenoidea 404 

Scorpaenoidei 401 

Scorpaenopsis cirrhosa 495 

Scutella perrini 297-298, 301, 303 

Scutellaster 298, 301, 357, 363, 369 
interlineatus 363-364, 369 
oregonensis 301, 369 
oregonensis quaylei 301 
sp. 364 

Searlesia sp. 364 

Siliqua patula 364 


Silvestriellus 427-428, 436, 443, 445, 448, 450-452 


flaviventris 427, 443-445, 449, 452 
martinezi 427, 443-445, 449, 452 
patagonicus 427, 443-445, 449, 452 


schlingeri 427, 436, 443-445, 449, 452 


sp. 445-446 
Simopias 475-476, 483-484 
didactylus 490 


Sindoscopus 125-126, 129-132, 136, 141-143, 145, 149- 


150, 158 


australis 126, 128, 131-132, 134, 136, 141-143, 


145, 148-149, 150-154, 158-159 

Sirembo 398 

grandis 390, 398 
Solen sicarius 363-364 
Sphagemacrurus 235-236, 238, 242-243 
Sphenomorphus 264 

brunneus 264 

cinereus 264 

nigrolineatus 264 

solomonis 264 
Spisula 

albaria 300 

albaria coosensis 364, 368-369 

mercedensis 364 

mossbeachensis 364 
Squalogadus 250 

modificatus 234, 250 
Squamata 523 
Squilla 484 
Stauroneis 

lauenburgiana 343 

montana 343 

montana f. lanceolata 343 
Stephanoberycidae 228, 230-231 
Stephanoberyciformes 227, 230 
Stephanoberycoidei 227 
Stephanodiscus 

alpinus 343 

astraea 343 

carconensis 343 

damasi 343 

dubius 343 

hantzschia 343 

lucens 343 


niagarae 343 
novaezeelandiae 343 
tenuis 343 
Sthenopus 404 
mollis 404—405 
Stylactis minoi 454, 458 
Stylemys 513 
Surirella 
amoena 343 
contorta 343 
engleri 343 
obscura 343 
skvortzowi 343 
sovereigni 343 
spinosa 343 
spiralis 343 
Swiftopecten swifti parmeleei 300 
Synanceia 480 
(Aploactis) aspera 404—405 
aspersa 419 
didactyla 489 
rubicunda 489 
Synanceidae 476—477 
Synanceiidae 453, 476 
Synancelinae 453 
Synanceinae 476 
Synanceioidei 476 
Synanceja 
didactyla 490 
rubicunda 490 
Synanciidae 476 
Synedra capitata 343 


Tabanidae 345, 427-428, 450-451 
Tabellaria 
binalis 343 
fenestrata var. asterionelloides 343 
Tamarix 521—522 
Testudinella horsfieldii 513 
Testudines 506 
Testudinidae 509, 511 
Testudo 501, 513-514, 516 
baluchiorum 513 
buxtoni 515 
caspica 511 
coriacea 509 
ecaudata 515-516 
euphraticus 519 
graeca 501, 514-516, 519, 525 
graeca graeca 516 


graeca ibera 501, 506, 513, 514-516, 518-519, 


§25-—526 
graeca terrestris 516, 525 


graeca zarudnyi 501, 506, 514-515, 516-519, 525— 


526 
hermanni 514 
horsfieldii 513-514 
ibera 513-514, 516, 518-519 


INDEX 


japonica 506 
kleinmanni 514, 516 
marginata 514 
orbicularis 510 
zarudnyi 515-516, 518 
Tetracyclus rupestris 343 
Tetraroge 
dermacanthus 416 
kagoshimensis 410, 403, 407, 416 
monacanthus 405 
Tetraroginae 404 
Trachonurus 235 
Transennella tantilla 364, 368-369 
Tresus 
nuttallii 363-364 
sp. 364 
Trichardis 187 
Trichopleura 404 
Triclis 192, 194, 196 
argentifacies 194, 196 
flavipes 196 
flavipilis 196, 199 
tagax 194, 196, 200, 202 
Trigla rubicunda 489-490 
Triglidae 477 
Trionychidae 519 
Trionyx 503, 519-520 
aegyptiacus 519 
euphraticus 501, 505, 519-520, 523 
ferox 520 
swinhoei 520 
triunguis 519 
Trogonophidae 502, 523-524 
Turbonilla sp. 364 


Umbelliflorae 371-372, 374-376, 383-385, 387 
Uropterygius 164, 167-168 

alboguttatus 167 

goslinei 161-162, 165, 167-168 


539 


Kamar 161-162, 164—168 
marmoratus 168 
xanthopterus 166-168 


Varanus griseus 521 
Ventrifossa 238, 240 
(Lucigadus) 238 


Xenoberyces 230 


Yoldia 

limatula 364 

scissurata subsp. strigata 300 
Yucca 386 

elata 386 

glauca 384-385 

rostrata 386 


Zabrops 183-186, 188, 194, 196, 198, 200, 209, 211 

flavipilis 183-184, 196, 198, 199-200, 204-205, 
208-209 

janiceae 183, 185, 198, 203-205, 208, 210, 211-212 

spp. 201, 203-205 

tagax 186, 196, 198, 200, 202, 206, 208 

tagax argutus 183, 185, 198, 200-201, 204-206, 
208 

tagax group 198, 200-201, 207, 210 

tagax spp. 200-201 

tagax tagax 183, 185, 189, 196-198, 200, 202, 204— 
206, 208 

thologaster 183, 185, 198, 203-205, 208, 210-212 

thologaster group 198, 203, 210 

wilcoxi 183, 198, 200, 206, 208-210 

wilcoxi arroyalis 183, 185, 198, 201, 204-205, 208- 
209 

wilcoxi playalis 183, 185, 198, 200-201, 204-205, 
207-209 

wilcoxi wilcoxi 183, 185, 198, 200-201, 204-205, 
206-209 


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